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ecology of phasmids - KLUEDO - Universität Kaiserslautern

ecology of phasmids - KLUEDO - Universität Kaiserslautern

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Life history & potential population growth 31<br />

3 Life cycle, potential population growth, and egg hatching<br />

failure <strong>of</strong> Metriophasma diocles<br />

3.1 Introduction<br />

Great variation in patterns <strong>of</strong> population densities clearly exists in herbivorous insects, but generally<br />

most insect species in the tropics occur in low numbers (e.g., Elton 1973; Basset et al. 1992; Basset<br />

1997, 1999; Novotny & Basset 2000). Such low population densities reflect the biotic potential <strong>of</strong> a<br />

species in the limiting boundaries <strong>of</strong> habitat capacity, intraspecific competition and regulating<br />

mechanisms within trophic cascades.<br />

A whole wealth <strong>of</strong> factors acts on individuals in a population thereby affecting net recruitment (i.e.,<br />

births minus deaths). These factors either do not respond to population density or are density-dependent<br />

and can be roughly categorized as: (1) intraspecific density-dependent factors, (2) environmental<br />

density-dependent factors, and (3) density-independent factors.<br />

The upper limit <strong>of</strong> population size is defined by intraspecific competition. Intraspecific competition as<br />

density-dependent regulation factor increases with population size. Higher competition leads to<br />

increasing death and decreasing birth rates resulting in a population decline (and vice versa). At a<br />

certain density, birth and death rate are equal and there is no net change in population size. This density<br />

is known as carrying capacity (K). The carrying capacity represents the population size the resources <strong>of</strong><br />

the environment can just maintain without a tendency to either increase or decrease (Begon et al. 1996).<br />

However, natural populations lack simple carrying capacities. Alongside with intraspecific competition,<br />

populations are regulated by multiple factors reducing the population size below the carrying capacity.<br />

Most factors in the biotic setting <strong>of</strong> a species respond to population size. Density <strong>of</strong> prey translates into<br />

predator densities (Volterra 1926; Lotka 1925), and diseases spread faster in higher populated areas<br />

(e.g., Stevenson 1959). Increasing population density also involves decreasing food availability and<br />

higher levels <strong>of</strong> interspecific competition (reviewed in Schoener 1983).<br />

Other biotic factors like intrinsic food quality or breeding sites on the other hand do not relate to<br />

population density, but their limited nutritious value or availability may act as regulators (Andrewartha<br />

& Birch 1954, 1984). Likewise, abiotic factors such as climate and natural disasters are density<br />

independent and can cause drastic impact on populations (e.g., Willig & Camilo 1991).<br />

Releasing a population from regulating constraints can uncover the impact <strong>of</strong> the above described<br />

control factors. When there are no limits on its growth, the population <strong>of</strong> a species will increase<br />

infinitely according to its biotic potential, i.e. the inherent power <strong>of</strong> an organism to reproduce and<br />

survive (Chapman 1931). The intrinsic rate <strong>of</strong> natural increase (r) will then be at its maximum. That<br />

means each individual <strong>of</strong> the population will contribute to population growth with peak reproduction<br />

(Begon et al. 1996). In natural populations, most individuals are not capable <strong>of</strong> peak productivity

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