Janzen–Connell hypothesis

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The Janzen–Connell hypothesis is a widely accepted explanation for the maintenance of tree species biodiversity in tropical rainforests. It was published independently in the early 1970s by Daniel Janzen[1] and Joseph Connell.[2] According to their hypothesis, there is a density- or distance-dependent factor in recruitment of seedlings from adults of tropical tree species due to host-specific predators or pathogens.[3] These predators/pathogens that specifically target a species make the areas directly surrounding that parent tree (the seed producing tree) inhospitable for the survival of seedlings. In other words, the hypothesis states that specialist predators, pests, and pathogens keep key plant species rare enough to reduce their competitive ability enough so as to make space available for many other species. A tree predator is an herbivore of some kind, often tropical insects, that consume parts of the tree for food. An example of a tree predator are the beetles in the Chrysomelidae family commonly referred to as Leaf Beetles. A tree pathogen refers to a disease caused by a bacteria, virus, or fungus that infects the tree. A tree parasite could also be classified as a pathogen or a predator of a species.

According to the Janzen–Connell hypothesis, those seedlings that are farthest from their parent have a competitive advantage as they do not suffer from the species-specific predators that are found more commonly around the parent. For example, the Black Cherry tree (Prunus serotina) seedlings farther from their parent tree are surrounded by fewer other cherry tree seedlings – in other words, they are less dense. Diseases and herbivores that prey on the Black Cherry tree are most commonly found and increase most rapidly in high-density cherry areas.[4] So, those cherry seedlings farther from the parent are less likely to encounter a Cherry tree parasite like Pythium and will more likely survive to adulthood in comparison to the cherry seedlings close to the parent.

This mechanism has been proposed as promoting diversity of forests as it promotes survival of a number of different plant species within one localized region. While previously thought to explain the high diversity of tropical forests in particular, subsequent research has demonstrated the applicability of the Janzen–Connell hypothesis in temperate settings as well. The Black Cherry is one such example of a temperate forest species whose growth patterns can still be explained by the Janzen–Connell hypothesis.

History[edit]

Daniel Janzen’s hypothesis[edit]

Daniel Janzen published his hypothesis in 1970 in The American Naturalist under the article "Herbivores and the Number of Tree Species in Tropical Forests."[1] His hypothesis was based on the observation that in tropical forests (when compared to temperate forests), there were few new adult trees in the immediate vicinity of their parent tree. He explained the low density of tropical trees and lack of "bunching" of tree types around parent trees for two reasons: (1) the number of seeds decline with distance from the parent tree and (2) that the adult tree, its seeds, and seedlings are a source of food for host-specific parasites and diseases. Using his observations, Janzen created a model demonstrating the probability of a seed maturation or a seedling survival as a function of distance from the parent tree (as well as total seed count, dispersal mechanism, and predatorial activity).

Joseph Connell hypothesis[edit]

Joseph Connell published his hypothesis in 1971 in Dynamics of Populations.[2] Unlike Janzen, Connell proposed experiments that focused on the key prediction that exclusion of host-specific predators would cause a decrease in diversity as tree species with greater establishment or competitive ability formed low-diversity seedling and sapling communities where dominance was concentrated in a few species.

He formed his hypothesis through observations in Queensland, Australia. Along with Jack Greening Tracey and Larry Johnson Webb, he mapped trees in two rainforests and observed that smaller seedlings tended to occur in single-species clumps. Smaller seedlings also exhibited greater mortality, especially when their nearest neighbor was an individual of the same species. This pattern lessened with growth and age until seedlings exhibited similar pattern diversity to adults. To reinforce these observations, Connell ran an experiment showing that adult trees have a deleterious effect on smaller trees of the same species. In another experiment, Connell found that pre-germination predation was greater on seeds near adults of the same species than those near adults of others. Through these observations, Connell suggests that each tree species has host-specific enemies that attack it and any of its offspring which are close to the parent. This emphasizes the importance of the role of predation in preventing trees from forming single-species groves, which is probably the only way in which one species of tree could exclude others by interspecies competition.

Effects on forest dynamics[edit]

Disease dynamics and tree density[edit]

Plant pathogens follow infectious disease dynamics. The basic reproductive rate (R_0) of a disease is dependent on three variables such that:


R0 = βL'S

Where β is the transmission rate or infectiousness of the disease, L is the average infection time of the host, and S is the density of the host population. By decreasing any one of the variables, the reproduction rate of the disease decreases. Since seed dispersal is such that the highest density of seeds is around the parent with density decreasing with distance from the parent, the reproduction rate of a disease infecting seeds and seedlings will be highest around the parent and decrease with distance. Thus, seedlings close to the parent are likely to die due to the disease prevalence. However, seedlings farther away are less likely to encounter the disease and therefore will more likely grow into adults.

Herbivory and tree density[edit]

Specialist herbivores who consume plant matter can also be thought of as having a "transmission rate" between individuals similarly to a disease. Tree predators (especially herbivorous insects) are limited by the ease of movement. When individuals are closer together at high density, movement between trees is easier and the predators quickly spread out. However, at low tree density, predators can not find the next individual with as much ease and thus often have low transmission rates leading to less specialist predation.

Concerns with the Janzen–Connell hypothesis[edit]

Many studies examining the Janzen–Connell hypothesis have shown supporting patterns with a number of tree species, but despite this there are also problematic aspects of the hypothesis.

  1. The first is that the Janzen–Connell hypothesis explains diversity at a community-wide scale, but most studies have only looked at a single species or a localized region. Most studies do not test the diversity prediction and did not determine the causes for patterns consistent with Janzen–Connell effects.[3] As Wright (2002) pointed out, "field measurements only demonstrate that niche differences, Janzen–Connell effects, and negative density dependence occur. Implications for species coexistence and plant diversity remain conjectural."[5]
  2. While these host-specific predators may play keystone species roles as they possibly aid in the prevention of superior tree competitors from monopolizing an area, no study has yet to examine species richness and abundance after removing natural enemies, creating a gap in the supporting research for the hypothesis.[3]
  3. Diversity may be maintained, at least in part, by episodic outbreaks of specialized pests, which may reduce the survivorship, growth, and reproductive success of adults of a species whenever they are particularly aggregated. Thus, specialist predation also impacts adult density, rather than just juveniles which has been the focus of research efforts.[3]
  4. Disease and predation may be just affecting overall density rather than in localized regions around adults alone.[3]

Research testing the hypothesis[edit]

Supporting research[edit]

There have been over 50 studies designed to test predictions of this hypothesis in tropical and other settings and most show that many tree species exhibit patterns consistent with Janzen–Connell effects. Studies that have supported the Janzen–Connell hypothesis:

  • A 2010 study of Panama forest reinforced that Janzen–Connell effects are also prevalent in tropical rainforests as seedlings growing close to their parents are likely to die due to microorganisms in the soil.[6]
  • A 2008 study by Petermann et al. in grasslands showed that soil-borne pests created a feedback very similar to the Janzen–Connell effect, supporting the hypothesis as a driver of diversity in temperate ecosystems. This study suggests that the predator/density mechanism that promotes species diversity is not contained to tropical forests alone – even if tropical ecosystems do have the highest diversity. This supports the hypothesis as a mechanism for diversity, but not as an explanation latitudinal gradients in species diversity.[7]
  • An examination of spatial data for 24 woody tropical rain forest plants showed either density-dependence or distance-dependence in plant offspring, supporting the hypothesis. However, other causal factors such as allelopathy were also suggested in some cases.[8]

Dissenting Research[edit]

Studies questioning the Janzen–Connell hypothesis:

  • Hyatt et al. examined the hypothesis across a number of tree species (rather than focusing on the effects on just one) and found no support for the distance-dependence of the hypothesis and concluded that the Janzen–Connell hypothesis was not a mechanism for diversity. They did find however that in temperate settings, distance from parent reduced survivorship. But in tropical settings there was a slight positive correlation between the improved competitive ability of seeds and distance from parent.[9]
  • A 1994 study by Burkey found that seed predation did not follow a pattern supporting that of the Janzen–Connell hypothesis along a meaningful scale. Seeds within 1 metre from the trunk were highly predated. However, seeds reached their peak density while still under the canopy of their parent. The authors concluded that seed predation did not follow the Janzen–Connell hypothesis.[10]

Current conclusions and further research[edit]

It is tricky to form conclusions regarding the accuracy of the Janzen–Connell hypothesis as it is difficult to falsify. This is because:

  • Heavy predation may keep some species rare and widely spaced, and these species may also be the best competitors. If so, these species are most likely to form dense aggregations that would reduce diversity in local areas. But due to their rarity or a belief that rare species are not regulated by density-dependence, these species may be the ones least likely to be studied.[3]
  • This means that the failure to find Janzen–Connell effects for what could be hundreds of tree species does not reject the hypothesis, as ecologists are missing the few key species where it does apply. However, dissenting findings do reduce the importance of the hypothesis for explaining overall coexistence of a number species.[3]
    • For example, Hyatt et al. (2003) found that there were "individual cases of conformity to the hypothesis,"[9] which is all that is needed for the hypothesis to work if the specific cases represent tree species that are excellent competitors, highly shade tolerant, habitat generalists, or some combination of these traits that would allow these species to otherwise dominate the ecosystem.[3]

It is likely that a number of mechanisms underscore the coexistence of similar species and thus cause biodiversity in ecosystems. It is possible the Janzen–Connell hypothesis is applicable only for some species depending on species characteristics. The hypothesis may also be affected by the kind of predator or pathogen as preliminary research has shown that the hypothesis is true only when host-specific predators have limited mobility with a range less than the seed dispersal range.[11]

References[edit]

  1. ^ a b Janzen, Daniel H. "Herbivores and the Number of Tree Species in Tropical Forests." The American Naturalist. 104.940 (1970).
  2. ^ a b Connell, J.H. "On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees." In: Dynamics of Population. Ed. P.J. Den Boer and G.R. Gradwell. Wageningen: Pudoc, 1970.
  3. ^ a b c d e f g h Carson, Walter A. and Stefan A. Schnitzer. Tropical Forest Community Ecology. Blackwell Publishing: Hoboken, NJ, 2008.
  4. ^ Packer, Alissa and Keith Clay. "Soil Pathogens and Spatial Patterns of Seedling Mortality in a Temperate Tree." Nature. 404. (2000): 278–281.
  5. ^ Wright, J.S. "Plant diversity in tropical forests: a review of mechanisms of species coexistence." Oecologia. 130.1 (2002): 1–14.
  6. ^ Mangan, Scott A., et al. "Negative plant–soil feedback predicts tree-species relative abundance in a tropical forest." Nature 466 (8 July 2010) 752–755.
  7. ^ Pertermann, Jana S. et al. "Janzen–Connell Effects are Widespread and Strong Enough to Maintain Diversity in Grasslands." Ecology 89.9 (2008): 2399–2406.
  8. ^ Clark, Deborah A. and David B. Clark. "Dynamics of a Tropical Rain Forest Tree: Evaluation of the Janzen–Connell Model." The American Naturalist. 124.6 (1984):769–788.
  9. ^ a b Hyatt, Lauren A. et al. "The Distance Dependence Prediction of the Janzen–Connell Hypothesis: a Meta-Analysis." Oikos. 103.3 (2003): 590–602.
  10. ^ Burkey, T.V. "Tree Species Diversity: A Test of the Janzen–Connell Model." Oecologia. 97. (1994): 533–540.
  11. ^ Nathan, Ran and Renato Casagrandi. "A simple mechanistic model of seed dispersal, predation and plant establishment: Janzen–Connell and beyond." Journal of Ecology. 92. (2004): 733–746.