connell's elegant field experiment wikipedia

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• 23 October 2020

Joseph H. Connell (1923–2020)

• Jane Lubchenco 0 &
• Wayne P. Sousa 1

Jane Lubchenco is a professor in the Department of Integrative Biology at Oregon State University in Corvallis. She spent a formative year in Connell’s lab in 1970–71.

You can also search for this author in PubMed   Google Scholar

Wayne P. Sousa is a professor of integrative biology at the University of California, Berkeley. He joined Connell’s lab in 1973, completing both a master’s and a PhD.

You have full access to this article via your institution.

Credit: Tad Theimer

Joseph (Joe) Connell altered both what and how ecologists study. Tree by tree, coral by coral, barnacle by barnacle, he saw patterns and processes across diverse ecosystems. Simply and with incontrovertible evidence, he demonstrated that interactions such as competition and predation could determine where species lived.

Before his classic experiments on Scotland’s rocky shores, field ecology was mainly descriptive, focusing on physical conditions such as temperature or moisture in determining where species lived. Connell, who died last month aged 96, inspired thousands of ecologists to test their hypotheses by manipulating conditions in the field.

Connell established long-term studies of coral reefs at Heron Island in the Great Barrier Reef and of tropical rainforests in Queensland, Australia, that spanned more than three and five decades, respectively. Monitoring revealed the dynamic nature of plant and animal communities that had long been considered stable. He discovered that natural variability in biological interactions and physical factors maintains diversity in these and other endangered ecosystems.

Born in 1923, Connell grew up just outside Pittsburgh, Pennsylvania. When the United States entered the Second World War in 1941, he enlisted in the Army Air Corps and was trained in meteorology. Later, conducting weather surveillance in the Azores — the Portuguese Atlantic archipelago — in support of army operations in Europe, he spent his free time birdwatching and identifying trees. Meeting army recruits who worked as wildlife managers, he realized it was possible to have a career as a biologist. After the war, and a degree in meteorology at the University of Chicago, Illinois, he headed to the University of California, Berkeley, for a master’s in zoology.

Connell produced what he described as a dull, unsatisfying thesis on brush rabbits ( Sylvilagus bachmani ) in the Berkeley Hills. Discouraged by the difficulties of conducting a population study (he trapped only 40 rabbits in 2 years), he adopted a rule of thumb — never again to study anything bigger than his thumb. As a doctoral student at the University of Glasgow, UK, he gleefully discovered what Charles Darwin had found a century before: that thousands of barnacles could easily be studied on the seashore, no traps required.

Connell realized that he could test his hypotheses about what factors determined where on the shore certain species lived by removing, adding or transplanting barnacles and their snail predators. Classic papers ensued, inspiring other ecologists to rethink distribution patterns, and, importantly, to test their ideas with controlled field experiments.

After a postdoc at the Woods Hole Oceanographic Institution in Massachusetts, Connell joined the faculty at the University of California, Santa Barbara, where he remained for the rest of his career. He was curious about processes that affected distribution and abundance, and those that might keep biodiversity high. Shifting to species that live for hundreds or thousands of years on coral reefs and in rainforests, he set up his Australian long-term monitoring studies in 1962 and 1963. Both recorded the demography and interactions of organisms in permanent plots, tracking community dynamics and the impact of disturbances, ranging from fallen trees to cyclones.

Visiting Connell’s sites with him in the 1970s and 1990s, we were impressed with his foresight and inspired by his insights. On the reef, he explained, physical disturbance by large waves associated with recurring cyclones intermittently reduced the cover of dominant species such as staghorn coral ( Acropora aspera ). This prompted recolonization by a diverse assemblage of weaker competitors such as encrusting or mound-like species. Connell coined the term ‘intermediate disturbance hypothesis’ to describe this process.

We strolled through the larger of his rainforest plots, avoiding stinging trees, biting flies, ticks and leeches, and relishing the richness — more than 300 tree species and about 100,000 individual plants. Connell outlined another hypothesis, that forests are more diverse when rarer species such as the conifer Sundacarpus amarus are favoured over more common ones such as the flowering tree Planchonella sp. Patterns of seedling establishment, growth or survival depend on that difference in frequency. Because common species grow more densely than rare ones, they are more vulnerable to specialist herbivores or pathogens.

This pattern of density-dependent predation or infection thins out common species, enabling a richer mix to coexist. It is a central component of the Janzen–Connell hypothesis (independently proposed by US ecologist Daniel Janzen in 1970), which predicts that seedlings are more likely to die under the canopies of their parent trees than farther away, ensuring diversity.

Connell was unfailingly kind, generous and devoted to his family. He never lost his profound curiosity about the natural world or his delight in exploring ideas with students and colleagues. He loved to be challenged and, if proven wrong, he gladly moved on to a new hypothesis or question. He sought truth, not fame. Moreover, he empowered everyone around him to think critically by focusing on ideas and evidence, not personalities. Fortunately for the world, his way of exploring science proved powerful, infectious, fun and enduringly productive.

Nature 586 , 670 (2020)

doi: https://doi.org/10.1038/d41586-020-02990-2

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Evidence for competition from nature Connells barnacles

The classic experimental demonstration of competition in the field was done by Joseph Connell (1961a, 1961b) on the barnacle species Chthamalus stellatus and Balanus balanoides. These two species are found growing in the rocky intertidal zone off the coast of Scotland. Intertidal zones frequently show vertical zonation of species based on their abilities to survive periods of exposure to the air during low tides, and wave action followed by submersion during high tides. Balanus is consistently found on lower rock surfaces, usually near mean tide level or slightly above. Chthamalus, however, is found on the upper rocks, between mean high neap tide and mean high spring tide. While the adults of these two barnacle species have non-overlapping distributions, the larvae of both species settle over a wide variety of rock surfaces, showing a great deal of overlap. The question Connell posed was, is the distribution of adults the result of competition, or is there a difference in the fundamental niches of the two species? Connell performed a variety of experiments in which he moved the barnacles to different levels of the intertidal zone. He also experimentally removed one species or the other where the two were growing together, and observed the results of putting the two species together. He found that whenever he removed Balanus, Chthamalus was able to survive in the lower regions of the intertidal zone. However, in the presence of Balanus, Chthamalus was overgrown and eventually displaced. In the upper regions of the intertidal zone, however, Balanus was unable to survive the long exposures to air during low tides. Since Chthamalus was able to survive this exposure, it survives in the upper intertidal zone. Thus the two species occupy mutually exclusive microhabitats due to a combination of competition and differences in their fundamental niches.

Direct observations of competition in ants

Because both worker and soldier ants are numerous, easy to observe, and usually diurnal, aggressive interactions among ant species, demonstrating interference competition, can be documented throughout the world (Holldobler and Wilson 1990). Placing a food bait of tuna or sugar water will provoke competitive interactions in a matter of minutes to hours. Once bait is put out in the West Indies, where there are few ant species, there is a kind of predictable sequence, reminiscent of ecological succession (a kind of "ant succession"). As described by Holldobler and Wilson (1990), first to arrive are workers of Paratrechina longicornis, known locally as "hormigas locas" (crazy ants). These workers are very adept at locating food and often are the first to arrive at newly placed baits. They fill their crops rapidly and hurry to recruit nestmates with odor trails laid from the rectal sac of the hindgut. But they are also very timid in the presence of competitors. As soon as more aggressive species begin to arrive in force, the Paratrechina withdraw and search for new, unoccupied baits. Paratrechina is an example of an "opportunist" species. They are poor competitors, but excellent dispersers. Next to arrive are species known as "extirpators." These species recruit other workers by odor trails and fight it out with competitor species. Examples include species in the genera Pheidole and Crematogaster, the fire ant (Solenopsis geminata), and the "little fire ant" (Wasmannia auropunctata). Some of these species have well-developed soldier castes that play a key role in the aggressive interactions. Injury and death are commonplace, and one species eventually dominates the bait. Pre-emption is usually the deciding factor. The colony whose foragers arrive first typically wins; foragers recruit nestmates, who surround the bait. When worker scouts encounter a large number of workers from another colony, they are easily repulsed (Holldobler and Wilson 1990). Species with a third strategy, called "insinuators," also arrive at the baits. These are small colonies with small-sized worker ants such as Tetramorium simillimum and species of Cardiocondyla. A scout who discovers the bait will recruit only one nestmate at a time. Small size and stealthy behavior allow these individuals to take some of the bait without provoking a response from the extirpator species, - a situation reminiscent of small animals sneaking in and removing bits of food at a lion kill.

Holldobler and Wilson also emphasize that territorial fighting and "ant wars" are common, especially among species with large colonies. Numerous cases have been documented in which introduced ant species have eliminated other species over a few years' time. For example, on Bermuda Iridomyrmex humilis has been replacing Pheidole megacephala since the former was introduced in 1953, although the two species may be reaching equilibrium short of extinction of Pheidole (Lieberburg et al. 1975). As a final example, the red imported fire ant (Solenopsis invicta) has virtually eliminated the native fire ant (S. xyloni) from most of its range in the United States (Holldobler and Wilson 1990).

Literature reviews of field studies on competition

In the 1980s the importance of interspecific competition in nature was questioned by a number of biologists. Strong et al. (1983), among others, challenged much of the evidence usually cited for the prominent role assigned to competitive interactions in structuring natural communities. They asserted that the data were often indistinguishable from random models. Others charged that there were few experimental studies of competition from nature, and that predation was a much more significant ecological interaction. Still others, such as Wiens (1977), asserted that competition is a temporally sporadic, often impotent, interaction. Schoener (1982, 1983) decided to review the literature to determine if competition had been affirmed as an important interaction in nature. He found, to his surprise, over 150 experimental field studies of competition in natural ("field" settings), many of which had been conducted in the previous five years. Schoener carefully defined an interspecific competition experiment as a manipulation of the abundance of one or more hypothetically competing species. All such experiments had to have proper controls. Prior to these experimental studies, there had been a dependence on "natural experiments," which will be discussed below (Diamond 1983).

The "field" was defined as a study in which some major natural factors extrinsic to the organism remain uncontrolled. Schoener did not allow laboratory or greenhouse setups, but did count experiments involving fenced exclosures or caged portions of shorelines to fit the definition of a field study.

Through 1982 Schoener found that 164 published studies fitted the criteria. Of those 164, 90% (148) of the studies and 76% of the species involved did show positive evidence of interspecific competition. In a separate analysis and using different criteria, Connell (1983) found evidence of competition in 40% of the experiments and 50% of the species. There were, however, few studies involving herbivorous insects that demonstrated interspecific competition. Schoener suggested, as had Hairston et al. (1960), that herbivores, which occupy an intermediate position in the food web, are controlled by predators and therefore competition is a less important interaction for this trophic position. Schoener's literature review found little evidence to support Wiens' idea that there is a great deal of temporal variability in competition. Competitive variability is especially rare in marine ecosystems. Variability was mostly found in dry, continental habitats. This is interesting in that Wiens developed his ideas after carrying out research on bird communities in North American arid or semi-arid shrub habitats.

A decade after the analyses of Schoener and Connell, Gurevitch et al. (1992) analyzed competition studies carried out from 1980 to 1989, using a statistical approach. They found "medium" effects of competition on primary producers, carnivores, and herbivorous marine mollusks. Larger effects of competition were detected on some herbivores and stream arthropods. As found by Schoener, however, studies on herbivorous terrestrial insects usually failed to show significant effects of competition.

One can conclude from these literature reviews that competition is a common event in nature that contributes to the organization of ecological communities. It is, however, not the only important interspecific interaction.

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Readers' Questions

What do you predict would happen if chthamalus were experimentally removed?

Why dont barnacle species overlap?

There are several reasons why barnacle species do not overlap: Competition for resources: Each barnacle species has specific requirements for survival, such as food availability, water temperature and salinity, substrate type, and wave exposure. If two barnacle species have similar ecological niches and requirements, they will compete for limited resources, leading to decreased fitness and population decline. Therefore, barnacle species tend to occupy different habitats where their specific needs are met, minimizing competition. Differential colonization abilities: Barnacles have limited dispersal abilities and rely on larval stages to settle and colonize new areas. Different barnacle species have varying abilities to colonize and establish populations in different habitats. Some species may have adaptations that make them more successful in intertidal zones, while others may be more successful in subtidal or offshore environments. This differential colonization ability contributes to the spatial separation of barnacle species. Biogeographical factors: Barnacle species distributions are also influenced by biogeographical factors such as ocean currents, geographical barriers, and historical events. These factors can restrict species dispersal and gene flow, leading to the formation of distinct populations and limiting the overlap of different species. For example, species on separate sides of a continental landmass or isolated islands may have little or no chance of overlap due to physical barriers. Ecological interactions: Barnacles interact with other organisms in their environment, including predators, competitors, and symbiotic species. These interactions can influence barnacle species distributions and overlap. For instance, if a particular predator species preys on one barnacle species, it may limit its population and restrict its overlap with other species that are not preyed upon. Similarly, if a barnacle species has a mutualistic relationship with a specific partner, they may be limited to areas where their partner species is present. Overall, the lack of overlap in barnacle species can be attributed to a combination of competition, differential colonization abilities, biogeographical factors, and ecological interactions.

How is the process of experimenting of connells barnacles with one?

The process of experimenting with Connell's barnacles involves several steps. Here is a general outline of the process: Selection of barnacles: Identify a population of barnacles to study. This could be done by collecting barnacles from a natural habitat or obtaining them from a supplier. Set up experimental tanks: Prepare tanks or containers filled with seawater to mimic the barnacles' natural environment. Ensure the tanks have appropriate lighting, temperature, and water quality. Acclimatize the barnacles: Gradually introduce the barnacles to the experimental tanks, allowing them to acclimate to the new conditions. This helps to reduce stress and ensure their survival during the experiment. Manipulate variables: Determine the specific variables you want to test. These could include factors like temperature, salinity, food availability, or water flow. Manipulate these variables in the experimental tanks to observe their effects on the barnacles. Control group: In a controlled experiment, it is essential to have a control group that is not subjected to the manipulated variable. This allows for comparison and helps determine the actual impact of the variable being tested. Observation and data collection: Regularly observe the barnacles and record relevant data. This may include growth rates, survival rates, reproductive success, or other measurable parameters. Use tools such as microscopes, calipers, or cameras to facilitate accurate measurements and observations. Replication: Repeat the experiment multiple times to ensure the results are consistent and reliable. This helps account for any variation or errors in the initial experiment. Analysis: Analyze the collected data using statistical methods to identify patterns, correlations, or significant differences between the experimental groups. Compare the results of the manipulated group(s) with the control group. Draw conclusions: Based on the analyzed data, draw conclusions about the effects of the manipulated variables on the barnacles. Determine if there were any observed trends, relationships, or impacts. Communication of results: It is crucial to communicate the findings of the experiment through scientific reports, presentations, or publications. This allows other researchers or interested parties to learn from the study and build upon it. Remember, specific protocols and equipment may vary depending on the complexity of the experiment and the research goals.

When was connell's barnacles and exclusion principle?

published Connell's barnacles and exclusion principle was published in 1955.

Why are controls not needed in the arthropods experiment described in the study guide?

Controls are not needed in the arthropods experiment because the study is focusing on the interactions between different species of arthropods and their environment. Since there is no manipulation of the environment or of the arthropods, there is no need to use a control group.

Can crazy ants conduct electricity?

No, crazy ants cannot conduct electricity.

Are barnacles herbivores?

No, barnacles are not herbivores; they are filter-feeding omnivores.

Which of the following compete for space on intertidal rocks?

• Mussels • Barnacles • Anemones • Seaweeds • Sea stars • Algae

Francis O'Leary's Working Title

Just another university of oregon sites site, the influence of interspecific competition and other factors on the distribution of the barnacle chthamalus stellatus. joseph connell (1961).

Upon witnessing that two species of barnacles seemed to have the adults of their species segregated into horizontal bands on a rocky intertidal shore but that the youth of the higher species could be found in the lower band with the other species, Connell set out to study the primary reason for that segregation. His theory was that the completion for space between the two species bore at least some responsibility for the separation. This theory was supported by the following studies; two species will either compete for resources with one species becoming more dominant in an area (Beauchamp and Ullyott 1932) (Kenny and Stevenson 1956), equal distribution of one species is due to that species competing primarily with itself (Holme 1950) (Clark and Evans 1954), and if two species with similar needs are living in the same area it is because they are not competing for resources (Lack 1954) (MacArthur 1958).

The method Connell used to test this theory was to map the locations of the barnacle species Chthamalus Stellatus, hereafter referred to at C.S. , in the period of the year before what he hypothesized to be C.S. ’s competitor Balanus Balanoides , hereafter referred to as B.B. . After mapping the locations of C.S. it was possible to control the height above or below mean tide level so that the effects of competition could be seen in environments where both C.S. and B.B. were primarily observed. One half of all clusters of C.S.  growth were kept from being interfered with by B.B. . The growth and mortality rates in each case were recorded.

The results showed that C.S. was fully capable of growing to maturity at the levels on which B.B. was typically dominant, implying that the competition for space is what was preventing C.S. ’s proliferation at the lower levels. In fact, while the hypothesis was that competition was at least somewhat responsible for the distribution of the two species, the study found that predation by carnivorous aquatic snails, battery by waves, and intraspecies crowding combined were not much more likely to be responsible for the death of an individual C.S. than was crowding of some sort by B.B. .

I read this essay about two hours ago and have been idly trying to come up with an example of competitors coexisting. None come to mind except in the case of lions and tigers and bears coexisting in Oz. Given this, I can’t find fault in Connell’s reasoning. His experiment took into account as many variables as I could think of and the information gained fulfilled his hypothesis without assumption. In the case of this essay the limited resource in question was space, but it is easy to imagine that if the same experiment were performed on species competing for water or a nutrient source like meat the results would show that at least some of the reason for animal dispersion was due to interspecific competition. As a follow up to this study, I would be interested to see if there were more studies done on intraspecific competition and what its causes may be.

15 thoughts on “ The Influence of Interspecific Competition and Other Factors on the Distribution of the Barnacle Chthamalus Stellatus. Joseph Connell (1961) ”

After mapping the sites of C.S. it was feasible to alter the height above or below mean using soundcloud downloader so that the impacts of competition could be seen in habitats where both C.S. and B.B. were primarily observed. One half of all clusters of C.S. growth were kept from being interfered with by B.B.. The growth and mortality rates in each case were documented.

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Your note on “The Influence of Interspecific Competition and Other Factors on the Distribution of the Barnacle Chthamalus Stellatus” by Joseph Connell provides a detailed and insightful summary of the essay. Here’s a breakdown of your note:

1. Background: You introduce the context of Connell’s study, where he observed the segregation of two barnacle species into horizontal bands on a rocky intertidal shore and aimed to understand the primary reason behind this segregation.

2. Theory and Support: You outline Connell’s theory that competition for space played a role in the separation of the barnacle species. You mention various studies that supported the idea that competition influenced species distribution.

3. Method: You describe the methodology Connell employed, which involved mapping the locations of Chthamalus Stellatus (C.S.) and controlling the height above or below mean tide level to study the effects of competition. Half of the C.S. clusters were protected from interference by Balanus Balanoides (B.B.).

4. Results: You summarize the key findings of the study, which showed that C.S. could grow to maturity at levels dominated by B.B., indicating that competition for space was a significant factor in the distribution of C.S.

5. Reflection: You reflect on the essay, noting that you couldn’t think of any examples of competitors coexisting, which supports Connell’s reasoning. You appreciate the thoroughness of Connell’s experiment and its consideration of various variables.

6. Interest in Further Studies: You express interest in further studies on intraspecific competition and its potential causes, suggesting that such research could provide valuable insights.

Overall, your note provides a well-structured summary of Connell’s study, highlighting its significance in understanding interspecific competition and species distribution. Your reflection and curiosity about further research demonstrate a strong engagement with the topic. Well done!

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Connell’s elegant field experiments on rocky sea coasts of Scotland is the evidence of

Habitat diversification

Prudent predators

Competitive release

Competitive exclusion

The correct option is D Competitive exclusion Competitive exclusion i.e. removal of inferior by a superior. Here Connell's Elegant field experiment showed that on rocky sea coasts of Scotland, the larger and superior barnacle Balanus dominates the intertidal area, and excludes the smaller barnacle Chathalamus from that zone.

Connell’s elegant field experiments on rocky sea coasts of Scotland is the evidence of

The steep rocky coasts rising almost vertically above the sea level are called

Connel's field experiment on the rocky sea coast of Scotland, where larger Barnacle balanus dominates the intertidal area and removes the smaller Barnacle cathamalus. This happened due to. Predation Competition Parasitism Mutualism

Connel's field experiment on the rocky sea coast of scotland, where larger barnacle balanus dominates the intertidal area and removes the smaller barnacle cathamalus. this happened due to competition for space between two species of barnacles, balanus balanoides and chthamalus stellatus. though chthamalus occupying an upper zone, and balanus, a lower zone. larvae of both species settle and attach over a wider vertical range than the zone occupied by adults. but barnacles are removed because chthamalus are more tolerant of physical desiccation than balanus. connell suggests that the lower limit of distribution of intertidal organisms is usually determined mainly by biotic factors, such as competition with other species and predation, whereas the upper limit is more often set by physical factors, such as dry conditions prevailing during low tides..

Connell’s elegant field experiments on rocky sea coasts of Scotland is the evidence of

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Structured data

Items portrayed in this file, copyright status, copyrighted, copyright license, creative commons attribution-sharealike 3.0 unported, source of file, original creation by uploader, coordinates of the point of view, 55°45'36.608"n, 37°39'10.822"e, exposure time, 0.0005 second, focal length, 29 millimetre, instance of.

• Naryshkiny estate (Maly Kazenny Lane)
• Moscow photographs taken on 2008-06-08
• CC-BY-SA-3.0
• Self-published work
• Files with coordinates missing SDC location of creation

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