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Population death in Hawaiian plant communities : a causal theory and its successional significance
(1983)
Population death or synchronized plant-group dying or dieback, as constrasted with individual plant dying or single plant death, is a common phenomenon in Hawaiian plant communities. Examples of population death are given from forest, scrub and grassland communities and from lowland, montane and higher altitude environments as well as from native and non-native plant populations.
In the better researched cases, the Metrosideros and Canavalia diebacks, it is clear that the dead standing populations are not simply relict stands left from herbivore predation of their reproduction. This explanation was given in the earlier literature for the dying Acacia koa and Sophora chrysophylla forest stands on Mauna Kea. Instead, in the Metrosideros and Canavalia diebacks and the other examples cited, reproduction from seedlings and/ or vegetative reproduction are typically associated with the dieback populations. The dieback populations themselves can be considered as cohorts, i.e. groups of individuals that originated from a wave regeneration in their respective habitats.
Five characteristics, common to the described dieback populations were noted: 1) the populations belong to pioneer or seral species, 2) they occur in synusiae with low species diversity, 3) they grow in areas where disturbances gave rise to invasion of large cohorts, 4) they are associated with reproduction of the same species under or nearby the dying cohort and thus were described as "oscillating persisters" in succession, and 5) the dieback populations persist in all cases for relatively long periods for several reasons (low rates of decomposition, slow rates of successional replacement and low frequency of catastrophic perturbations).
Current hypotheses to explain population death in plant communities were reviewed as emphasizing one of four possibilities: 1) diseases or biotic stresses due to fungal pathogens or insect pests, 2) new man-imposed stresses, 3) recurring abiotic natural stresses, and 4) combinations of stresses.
A new theory is proposed which takes all the described dieback variations into consideration. It focuses on the dieback events as a chain reaction process involving: 1) senescing cohorts as the major predisposing condition, 2) dieback precipitating or triggering factors. These may operate as species-specific internal triggers (such as a heavy flowering season in the senescing stage) or as hard-to-detect environmental triggers, for example, a strong local wind that may tear off much of the foliar biomass (which then cannot be replaced because of the low carbohydrate reserves in the senescing stage), and 3) dieback-hastening factors, such as biotic agents and/or also dieback-stalling factors such as a temporary more favorable soil water or nutrient condition.
In addition, this theory is seen as providing fresh insights into the successional consequences of such diebacks. Dieback may be considered a driving force in secondary succession whenever it occurs, because of the relatively sudden opening of the canopy or death of the shoot systems. This in turn releases nutrients through death of the root systems and high litter in puts and protects the surviving undergrowth species and new seedlings (or vegetative reproduction of the dying cohort) from competition. Moreover, several indications in the Hawaiian Metrosideros rain forest have led to the hypothesis that the next generation of Metrosideros seedlings is not always genetically and physiologically identical to the dying cohort on the same site. That is, there are successional races or successional ecotypes indicating that Metrosideros polymorpha may have evolved into its own successional replacer. This is seen as an analogy to floristically richer areas, where the successional replacers are usually different species which form a functional sequence from pioneer, seral to climax along successional gradients. The hypothesis of successional races or ecotypes in Metrosideros polymorpha is currently subjected to experimental research in Hawaii using the "Hohenheimer" water table model as a transplant garden.