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Revisiting Paine’s 1966 sea star removal experiment, the most-cited empirical article in the American Naturalist

“Food Web Complexity and Species Diversity” (Paine 1966) is the most-cited empirical article published in the American Naturalist . In short, Paine removed predatory sea stars ( Pisaster ochraceus ) from the rocky intertidal and watched the key prey species, mussels ( Mytilus californianus ), crowd out seven subordinate primary space-holding species. However, because these mussels are a foundational species, they provide three-dimensional habitat for over 300 associated species inhabiting the mussel beds; thus, removing sea stars significantly increases community-wide diversity. In any case, most ecologists cite Paine (1966) to support a statement that predators increase diversity by interfering with competition. Although detractors remained skeptical of top-down effects and keystone concepts, the paradigm that predation increases diversity spread. By 1991, “Food Web Complexity and Species Diversity” was considered a classic ecological paper, and after 50 years it continues to influence ecological theory and conservation biology.

American Naturalist

Walden Pond, Massachusetts: Environmental setting and current investigations

Introduction Walden Pond, in Concord, Massachusetts, is famous among lakes because of its unique social history. Walden was the setting for American naturalist Henry David Thoreau's well-known essay 'Walden; or, Life in the Woods,' first published in 1854. Thoreau lived and wrote at Walden Pond from July 1845 to September 1847. In 'Walden,' Thoreau combined highly admired writing on Transcendental philosophy with pioneering observations of aquatic ecology and physical aspects of limnology, the study of lakes. Because Thoreau also defended so effectively the value of living close to nature in the Walden woods, the pond is considered by many to be the birthplace of the American conservation movement. Visitors come from all over the world to the pond, which has been designated a National Historic Landmark, and its fame has resulted in a major fund drive to preserve the surrounding woods. Walden Pond has no surfacewater inflow or outflow, and much of its ground-water contributing area likely is preserved within the Walden Pond Reservation area (fig. 1). Only 15 miles from Boston, the pond is unusually clear and pristine for an urban-area lake. However, point sources of nutrients near the pond, and a large annual visitor attendance, concentrated during the summer when the swimming beach (fig. 2) is open, may contribute a nutrient load sufficient to change the pond environment. The occurrence of nuisance algal species, a recent beach closing, and an awareness of water-quality problems suffered by other ponds in the region raise concerns about the risk of ecological change at Walden Pond. Despite the role of Walden Pond as a cultural and environmental icon, little is known about the pond's ecological features, such as its internal nutrient cycling or the structure of its food web, nor have consistent measurements been made to determine whether these features are changing or are stable. Production rates of aquatic plants in lakes and ponds naturally undergo a slow increase as plant nutrients, organic matter, and soil are leached and eroded from the surrounding watershed. This process is known as eutrophication. The term 'cultural eutrophication' refers to an accelerated form of the natural process in which extra soil and nutrients are derived from people's use of fertilizer, rerouting of surface drainage, and disposal of domestic and industrial waste. Cultural eutrophication can lead to excessive growth of aquatic plants, pond filling by decayed plants and eroded soils, reduced water clarity, and depletion of dissolved oxygen in deep water with subsequent loss of cold-water fish habitat. In order to document the longterm ecological health of Walden Pond, the U.S. Geological Survey (USGS), working in cooperation with the Massachusetts Department of Environmental Management (MDEM), is investigating factors that could contribute to cultural eutrophication of Walden. Through measurements of mass balance of nutrients and oxygen in the pond's deep water, the investigation will establish a baseline data set on Walden's trophic state, which is a measure of the pond's ability to support plant growth. The baseline data will be used to detect trends and give early warning of trophic changes or trophic response to pond remediation projects. This Fact Sheet provides background information on the environmental setting, limnological features, and cultural eutrophication of Walden Pond, and describes the joint USGS/MDEM study.

Massachusetts

Latitudinal and photic effects on diel foraging and predation risk in freshwater pelagic ecosystems

1. Clark & Levy ( American Naturalist , 131 , 1988, 271–290) described an antipredation window for smaller planktivorous fish during crepuscular periods when light permits feeding on zooplankton, but limits visual detection by piscivores. Yet, how the window is influenced by the interaction between light regime, turbidity and cloud cover over a broad latitudinal gradi- ent remains unexplored. 2. We evaluated how latitudinal and seasonal shifts in diel light regimes alter the foraging- risk environment for visually feeding planktivores and piscivores across a natural range of turbidities and cloud covers. Pairing a model of aquatic visual feeding with a model of sun and moon illuminance, we estimated foraging rates of an idealized planktivore and piscivore over depth and time across factorial combinations of latitude (0 – 70 ° ), turbidity (0  1 – 5 NTU) and cloud cover (clear to overcast skies) during the summer solstice and autumnal equinox. We evaluated the foraging-risk environment based on changes in the magnitude, duration and peak timing of the antipredation window. 3. The model scenarios generated up to 10-fold shifts in magnitude, 24-fold shifts in duration and 5  5-h shifts in timing of the peak antipredation window. The size of the window increased with latitude. This pattern was strongest during the solstice. In clear water at low turbidity (0  1 – 0  5 NTU), peaks in the magnitude and duration of the window formed at 57 – 60 ° latitude, before falling to near zero as surface waters became saturated with light under a midnight sun and clear skies at latitudes near 70 ° . Overcast dampened the midnight sun enough to allow larger windows to form in clear water at high latitudes. Conversely, at turbidities ≥ 2 NTU, greater reductions in the visual range of piscivores than planktivores created a window for long periods at high latitudes. Latitudinal dependencies were essentially lost during the equinox, indicating a progressive compression of the window from early summer into autumn. 4. Model results show that diel-seasonal foraging and predation risk in freshwater pelagic ecosystems changes considerably with latitude, turbidity and cloud cover. These changes alter the structure of pelagic predator – prey interactions, and in turn, the broader role of pelagic consumers in habitat coupling in lakes.

Journal of Animal Ecology

Shippingport, Kentucky, is the type locality for the white-footed mouse, Peromyscus leucopus (Rafinesque, 1818) (Mammalia: Rodentia: Cricetidae)

The white-footed mouse, Musculus leucopus Rafinesque, 1818 (= Peromyscus leucopus), is a common small mammal that is widespread in the eastern and central United States. Its abundance in many habitats renders it ecologically important, and its status as a reservoir for hantavirus and Lyme disease gives the species medical and economic significance. The recognition of two cytotypes and up to 17 morphological subspecies of P. leucopus indicates considerable variation in the species, and to understand this variation, it is important that the nominate subspecies be adequately defined so as to act as a standard for comparison. Relevant to this standard for the white-footed mouse is its type locality, which has generally been accepted to be either the vague "pine barrens of Kentucky" or the mouth of the Ohio River. Newly assembled information regarding the life and travels of Constantine S. Rafinesque, the North American naturalist who described P. leucopus , establishes that Rafinesque observed this species in July 1818 while visiting Shippingport, Kentucky, which is now within the city limits of Louisville, Jefferson Co., Kentucky. Shippingport is therefore the actual type locality for this species.

Proceedings of the Biological Society of Washingto

Pranked by Audubon: Constantine S. Rafinesque's description of John James Audubon's imaginary Kentucky mammals

The North American naturalist Constantine S. Rafinesque spent much of the year 1818 engaged in a solo journey down the Ohio River Valley to explore parts of what was then the western United States. Along the way, he visited a number of fellow naturalists, and he spent more than a week at the Henderson, Kentucky, home of artist and ornithologist John James Audubon. During the succeeding two years, Rafinesque published descriptions of new species that resulted from his expedition, including eleven species of fishes that eventually proved to have been invented by Audubon as a prank on the credulous naturalist. Less well known are a number of “wild rats” described by Rafinesque that include one recognized species ( Musculus leucopus ) and ten other, imaginary “species” fabricated by Audubon ( Gerbillus leonurus , G. megalops , Spalax trivittata , Cricetus fasciatus , Sorex cerulescens , S. melanotis , Musculus nigricans , Lemmus albovittatus , L. talpoides , Sciurus ruber ). Rafinesque's unpublished sketches of these animals provide important insight regarding the supposed nature of the animals invented by Audubon and ultimately published by Rafinesque.

Archives of Natural History

Bat mortality and activity at a Northern Iowa wind resource area

We examined bat collision mortality, activity and species composition at an 89-turbine wind resource area in farmland of north-central Iowa from mid-Apr. to mid-Dec., 2003 and mid-Mar. to mid-Dec., 2004. We found 30 bats beneath turbines on cleared ground and gravel access areas in 2003 and 45 bats in 2004. After adjusting for search probability, search efficiency and scavenging rate, we estimated total bat mortality at 396 ?? 72 (95 ci) in 2003 and 636 ?? 112 (95 ci) in 2004. Although carcasses were mostly migratory tree bats, we found a considerable proportion of little brown bats (Myotis lucifugus). We recorded 1465 bat echolocation call files at turbine sites ( 34.88 call files/detector-night) and 1536 bat call files at adjacent non-turbine sites ( 36.57 call files/detector-night). Bat activity did not differ significantly between turbine and non-turbine sites. A large proportion of recorded call files were made by Myotis sp. but this may be because we detected activity at ground level only. There was no relationship between types of turbine lights and either collision mortality or echolocation activity. The highest levels of bat echolocation activity and collision mortality were recorded during Jul. and Aug. during the autumn dispersal and migration period. The fatality rates for bats in general and little brown bats in particular were higher at the Top of Iowa Wind Resource Area than at other, comparable studies in the region. Future efforts to study behavior of bats in flight around turbines as well as cumulative impact studies should not ignore non-tree dwelling bats, generally regarded as minimally affected. ?? 2011, American Midland Naturalist.

American Midland Naturalist

T.D.A. Cockerell (1866–1948) of the University of Colorado: His contributions to the natural history of the California islands and the establishment of Channel Islands National Monument

Theodore Dru Alison Cockerell was a naturalist at the University of Colorado from 1904 to 1947 and studied botany, zoology, and paleontology in North and South America, Asia, Australia, Africa, and Europe. In the latter part of his career, he studied the California islands and published many papers on their natural history, 16 of them in four years (1937–1940). He made important contributions to the natural history of the islands in four distinct ways: entomology of the islands, including identification of a number of new species of bees; discovery of fossil marine invertebrate faunas and recognition that the zoogeography of the taxa reflects the position of the islands relative to converging cool and warm currents; discovery of abundant land snails, both living and in fossil form, and recognition that the fossils are in close stratigraphic association with mammoth fossils; and island biogeography, with considerations of species dispersal mechanisms and endemism. Newly discovered letters and memos also reveal that Cockerell played a pivotal role in the establishment of Channel Islands National Monument.

Western North American Naturalist

Alcoholic fermentation in swamp and upland populations of Nyssa sylvatica: Temporal changes in adaptive strategy

Both the upland and swamp varieties of Nyssa sylvatica respond initially to flooding with an acceleration of ethanol production. Under continued flooding the roots of the upland variety sylvatica have decreased rates of ethanol production, very likely a result of the progressively worsening necrosis of the root system. Few of these plants survive a full year of flooding. The swamp variety biflora survives and grows well under flooded conditions due to sequential metabolic and anatomical changes in the roots. Within a month after flooding, var. biflora initiates new roots with greatly accelerated rates of ethanol production, perhaps providing for a short-term compensatory energy source. Long-term acclimation to flooding involves a replacement of these roots and increased oxygen transport to the roots. The hypoxic conditions of the soil environment are avoided, and these rates of ethanol production are similar to those of drained plants.

American Naturalist

On the variability of alligator sex ratios

Samples of alligators from wild and 'farm' populations exhibited disproportionate sex ratios. Males predominated among young alligators from wild populations, whereas females were much more abundant than males in the farm population, where resources were superabundant. These results and other considerations lead us to hypothesize that environmental factors influence sex determination in alligators. During favorable environmental conditions natural selection is expected to favor a preponderance of the sex whose individuals exhibit the greater environmentally associated variation in relative fitness. We hypothesize that environmentally associated variation in age at sexual maturity of females produces sufficient variation in relative fitness of females to result in selection for low sex ratios during periods of resource abundance.

American Naturalist

Optimal timing in biological processes

A general approach for obtaining solutions to a class of biological optimization problems is provided. The general problem is one of determining the appropriate time to take some action, when the action can be taken only once during some finite time frame. The approach can also be extended to cover a number of other problems involving animal choice (e.g., mate selection, habitat selection). Returns (assumed to index fitness) are treated as random variables with time-specific distributions, and can be either observable or unobservable at the time action is taken. In the case of unobservable returns, the organism is assumed to base decisions on some ancillary variable that is associated with returns. Optimal policies are derived for both situations and their properties are discussed. Various extensions are also considered, including objective functions based on functions of returns other than the mean, nonmonotonic relationships between the observable variable and returns; possible death of the organism before action is taken; and discounting of future returns. A general feature of the optimal solutions for many of these problems is that an organism should be very selective (i.e., should act only when returns or expected returns are relatively high) at the beginning of the time frame and should become less and less selective as time progresses. An example of the application of optimal timing to a problem involving the timing of bird migration is discussed, and a number of other examples for which the approach is applicable are described.

American Naturalist

Hotshots, hotspots, and female preference in the organization of lek mating systems

We critically review the female-preference and hotspot models, the two most widely accepted recent explanations of lek organization. On the basis of what we believe are the inadequacies of these models-too great a reliance on the presumed acuity of female discrimination, the assumption that females have full freedom of choice within the lek, and insufficient recognition of the importance of male-male interactions-we develop an alternative set of hypotheses, which we call the hotshot model, to explain the development and maintenance of lek behavior. Our model attributes strong male mating skew to the interaction between (1) simplified and conservative mating rules of females and (2) social dominance among males. We demonstrate the importance of male-male dominance relationships in lek and non-lek court mating systems. We then argue that a strong mating skew among males forces novice males entering a population to adopt a long-term mating strategy that involves delayed breeding (floating) and subordinate lek behavior. The structure of leks is created by a complex of malemale interactions, with conflict between hotshots (who attempt to control lek mating) and subordinates, who may benefit from disrupting lek activities. Explanations for the number of males in an arena and inter-arena distances are based on modifications of the hotspot and female-preference models. We suggest specific field tests to help distinguish which hypothesis best models the behavioral interactions that produce lek mating.

American Naturalist

Climatic control of vegetation distribution: The role of the water balance

The water balance describes climate as it is sensed by plants, as the interaction of energy and water in the environment. Discriminant analysis showed that the distribution of North American plant formations was more highly correlated with the water balance (actual evapotranspiration and deficit) than with the more traditional measures of climate (such as temperature and precipitation) used in several studies, including those used in the well-known works of Thornthwaite, Holdridge, and Whittaker. Much of the improved correlation could be attributed to the ability of the water balance to distinguish between climates similar in mean annual energy and water supplies but different in the seasonal timing of the two. Consideration of the water balance aided in the interpretation of possible mechanisms controlling the distribution of plant formations. For example, coniferous forest occurred at low actual evapotranspiration (low simultaneous availability of energy and water), consistent with the suggestion that conifers are better adapted than deciduous trees to environments with a low potential for primary production. A better understanding of the mechanisms by which climate controls vegetation distribution will help us predict the effects of changing climate on the future distribution of vegetation types.

American Naturalist

Survival rates of birds of tropical and temperate forests: Will the dogma survive?

Survival rates of tropical forest birds are widely assumed to be high relative to the survival rates of temperate forest birds. Much life-history theory is based on this assumption despite the lack of empirical data to support it. We provide the first detailed comparison of survival rates of tropical and temperate forest birds based on extensive data bases and modern capture-recapture models. We find no support for the conventional wisdom. Because clutch size is only one component of reproductive rate, the frequently assumed, simple association between clutch size and adult survival rates should not necessarily be expected. Our results emphasize the need to consider components of fecundity in addition to clutch size when comparing the life histories of tropical and temperate birds and suggest similar considerations in the development of vertebrate life-history theory.

American Naturalist

A structural equation model of plant species richness and its application to a coastal wetland

Studies of plant species richness have often emphasized the role of either community biomass (as an indicator of density effects) or abiotic factors. In this article we present a general model that simultaneously examines the relative importance of abiotic and density effects. General and specific models were developed to examine the importance of abiotic conditions, disturbance, and community biomass on plant species richness. Models were evaluated using structural equation modeling based on data from 190 plots across a coastal marsh landscape. The accepted model was found to explain 45% of the observed variation in richness, 75% of biomass, and 65% of light penetration. Model results indicate that abiotic conditions have both direct effects on the species pool and indirect effects on richness mediated through effects on biomass and shading. Effects of disturbance were found to be indirect via biomass. Strong density effects on richness were indicated by the results, and canopy light penetration was found to be a better predictor of richness than was biomass. Overall, it appears that richness in this coastal landscape is controlled in roughly equal proportions by abiotic influences on the species pool and density effects, with disturbance playing a lesser role. The structure of the general model presented should be applicable to a wide variety of herbaceous plant communities.

Louisiana, Mississippi