r/K Strategists concept Alan Soffan In nature insect makes adaptation known as adaptive stategy as a result of selective environment factor acting in a population (Horn, 1976). They need to have certain strategies for efficiently utilizing the available time and energy to maximize their own survival (Atkins, 1980).There are two extreme strategist applied by insect in adapting theirself in certain environment. 1. R-stretegies R-strategies principally operates to maximize reproductive rates in harsh, unstable environment or during habitat disturbance (Atkins, 1980; Matthews&Matthews, 1978), while there are only a little competition exist (Elzinga, 1997). This R-strategies favor a “gambling” strategies (Price,1984). Some characters for r-strategist listed in table below. 2. K-strategy Principally, these insect tend to maintain their populations at a more or less constant level, close to the carrying capacity of environment (Atkins, 1980), They prefer not to increase a number of young, but they apply their high survival ability (Matthews&Matthews, 1978), Some characters for r-strategist listed in table below.. Table: Characteristics of extremes of r-K strategist R-Strategist Body size small K-Strategist Body size large Short generation time Long generation time High level of dispersal Low level of dispersal The mortality is densityindependent High reproductive rate and often reach overshoot The mortality is density dependent Low reproductive rate and rarely reach overshoot Population density fluctuative “boom and bust” Tend to be poor competitors Good colonization ability Population density relative constant from generation to generation Good competitors or keen No food spezialitation Usually they restrict just to The population only contains a few of individuals Reference Matthews& Kitching, 1984; Horn, 1976 Matthews&Matthews, 1978; Matthews& Kitching, 1984; Horn, 1976 Matthews& Kitching, 1984 Matthews& Kitching, 1984; Horn, 1976 Matthews&Matthews, 1978; Matthews& Kitching, 1984; Horn, 1976 Matthews&Matthews, 1978; Matthews& Kitching, 1984 Matthews&Matthews, 1978; Horn, 1976 Matthews&Matthews, 1978; Matthews& Kitching, 1984 Matthews& Kitching, They lives in unpredictable environment Rarely able to persist successfully for long period in certain habitat Able to quickly discover new habitats or nomadens Examples: grasshoppers (Chortoicetes terminifera), wasp scelio fulgidus, Bushfly Musca vestustissima, few of plant They usually lives in a stable environment High survival rate, especially reproductive stages Examples : Morabine grasshoppers (Geckomima sp.) 1984 Matthews&Matthews, 1978; Matthews& Kitching, 1984; Horn, 1976 Matthews& Kitching, 1984 Matthews&Matthews, 1978; Matthews& Kitching, 1984 (Matthews& Kitching, 1984) Reference Atkins, Michael D.1980. Introduction to Insect Behavior.Mac Millan Publishing Co.New York.237p Elzinga, Richard J. 1997. Fundamentals of Entomology. Prentice Hall. New Jersey.475p. Horn, David J. 1976. Insect Biology.W.B.Saunders.Philladelphia.439p Matthews, Robert W., &Matthews, J.R. 1978. Insect Behaviour. John Wiley & Sons. Canada.507p. Matthews, E.G.& Kitching, R.L. 1984. Insect Ecology. University Quesland Press.Asutralia.211p. Price, Peter W.1984. Insect Ecology.John Wiley&Sons. New York.607p. By Arya Insects commonly have two type of strategy applied in its life. These strategies are r and K-strategy. Some of insect may be flexible in applying r-K strategy. Locust and aphids are example for r selected insect and tsetse flies and carpenter bees are example for k selected insect (Matthew and Kitching 1984).Flour beetles, Tribolium castaneum and Tribolium confusum, are best-studied example for insect that are flexible in applying r-K strategy.(Matthew, 1976) r-strategist occupy to environment which has high food density and low population density of insect (Speight et al 1999). Such situation may occur when there are changes in weather, or through human activity such as the planting of food crop. r strategists are likely to be unpredictable in their ecologies, to produce large number of offspring in relatively brief period of time, and essentially to over exploit their habitats (Matthews and Kitchin 1984, Robinson 1987). r strategist has high mobility and polyphagy (Matthew, 1976). They are very efficient at exploiting a seasonal food supply that is available for only short period of time (Robinson 1987). K strategist, on the other hand, are more often regulated by density-dependent factors, such as intraspecific competition and natural enemies, they have relatively few offspring and avoid over exploiting their habitats. Some species are characterized by having giant eggs, low fecundity, and prolonged life spans (Maeta et al.1992). they are also food specialization, individual defence, and lack of mobility (Matthew, 1976). R or k strategist have each own advantages. R strategist relies on number of individuals, while a K strategist relies on the quality of individuals. R strategist has a great flexibility at the system level of the population. This means that the species can survive major catastrophes, even if only at the expense of very large number of individuals. For example, in the presence of the insecticide DDT, a new DDT resistant strain can be selected, bur this can only happen at the cost of very high mortality (Robinson 1987). On the other hand, K strategist have great flexibility at the level of individual. They depend on the survival ability of a complex and resilient individual which is able to cope with all but the worst of natural catastrophes (Robinson 1987). In the nature, r strategist are controlled by density independent control. Study on the population of Euphydryas editha, shows that interspecific competition and predation play relatively small roles in controlling many (but not all) of its populations (Daly 1978). In contrast, a butterfly of humid tropics such as Heliconius ethilla, which said to be K selected, (Ehrlich and Gilbert 1973) is not subject to catastrophic weatherrelated mortality, but it faces heavy loses in the larval stage from predatory ants (Daly 1978). The best studied of flexible r-K strategy are Tribolium castaneum and Tribolium confusum (King and Dawson 1972). These beetles have intrinsic density-regulating mechanism, including egg and pupal cannibalism by larvae and adult, reduced fecundity and fertility in aged population, and the secretion of an inhibitory quinone gas in crowded situation (Matthew 1976). Cannibalism is not only a regulatory mechanism but also improves the fitness of surviving individuals (Mertz and Robert 1970). References 1. Daly, H.V. 1978. Introduction to insect biology and diversity. London: McGraw-Hill. 2. Matthews, E.G. 1976. Insect ecology. St Lucia: University of Queensland Press. 3. Matthews, E.G. and R.L. Kitching. 1987. Insect ecology 2nd edition. St Lucia: University of Queensland Press. 4. Robinson, R.A. 1987. Host management in crop pathosystems. New York: MacMillan Publishing Company. 5. Speight, M.R., M.D. Hunter, and A.D. Watt. 1999. Ecology of insect: concept and application. Oxford: Blackwell Science. r/K Strategists concept Boy r/k strategist is a life history theory proposed by Mac Arthur and Wilson (1967) for classifying species based on their population growth which has correlation with their life strategies to survive in the environment (Denno and Dingle, 1981; Robinson, 1987). In this concept, species are classified as r-strategists and Kstrategists (Zadoks and Schein, 1979; Pedigo, 1989). Insects (species) which have high reproductive rates but with low survival rates are called r-strategists, named after the statistical parameter “r”; the symbol for the growth rate coefficient (Pedigo, 1989). Many pests of crops are rstrategists, they are small in size, have high reproductive rate, have short-lived individuals in a relatively brief period of time, have high level of dispersal ( Robinson, 1987; Matthews and Kitching, 1984), have low investment in defence and other interspecific competitive mechanisms (Matthews and Kitching, 1984), and they have capacity to escape from severe enemy impact through the interaction of excellent colonizing ability and a high population growth (Metcalf and Luckmann, 1982). For instance, their colonizing ability can be seen from the explosion of the pest population in early season (after germination of the host plant) or after the application of insecticides, where the big number of new resistant population will be present for replacing the susceptible population (Metcalf and Luckmann, 1982). Migratory locust is an example of r-strategists. It was so difficult to control this pest which has ability to reproduce new generation rapidly (Matthews and Kitching, 1984) and has ability to disperse or migrate in a great distances (Metcalf and Luckmann, 1982). Water beetle (Eretes australis), ghost moths (Trictena argentata) and some of dragonflies are other examples of r-strategists species (Matthews and Kitching, 1984) In contrast, insects which have low reproductive rates but with high survival rates are called K-strategists, named after the symbol for flattened portion of a population growth curve “K”, the abbreviation means the carrying capacity of the environment (Pedigo, 1989; Robinson, 1987). K-strategists has characteristics such as large in size, low level of dispersal organisms, long-lived individuals over relative long periods of time (Robinson, 1987). K-strategists live in harsh environment by producing few offspring which have ability to defense from predation (Metcalf and Luckmann, 1982; Zadoks and Schein). Morabines grasshopper is an example of K-strategists. A Population of this species may contain only a few of individuals (low reproduction rate) but with the ability to survive in the environment by using mimicry/ camouflage on the plants where they are life (Matthews and Kitching, 1984). Tsetse flies, carpenter bees, and some forest lepidoptera are examples of K-strategists species (Speight et.al, 1999). We may find many intermediate species (between r and K strategiests) in the real world, and some species may show traits of both r- and K-strategists at different times or in different locations (Speight et.al, 1999). A single species may exhibit r- or K-strategists depending on its population density. Low population densities of the bean weevil (Acanthoscelides obtectus) showed r -strategists characteristics when compared with high-density populations, which appeared to be more K-strategists (Aleksic et al, 1993 in Speight et.al, 1999). REFERENCES Denno, R.F and Dingle, H. 1981. Insect Life History Patterns: Habitat and Geographic Variation. Springer-Verlag: New York. Matthews, E.G and Kitching, R.L. 1984. Insect Ecology. Second Edition. University of Queensland Press: Australia. Metcalf, R.L and Luckmann, W.H. 1982. Introduction to Insect Pest Management. Second Edition. John Wiley and Sons: New York. Pedigo, L.P. 1989. Entomology and Pest Management. MacMillan Publishing Company: New York Robinson, R.A. 1987. Host Management in Crop Pathosystems. MacMillan Publishing Company: New York Speight, M.R; Hunter, M.D; Watt, A.D. 1999. Ecology of Insects. Blackwell Science: London. Zadoks, J.C. and Schein, R.D. 1979. Epidemiology and Plant Disease Management. Oxford University Press: New York. By Haris Insects as another animals, has behavior to response the habitat or environment changes. Response to environment change and environmental changes produced the dynamical of insect it’s self. The dynamical population of insect known as r-/K- selection (another scientist called r-/K- strategist). r is symbol of rate of population growth and K is symbol population density. The number of individuals of species per unit area of space is referred to as population density, and growth of population is the number of individuals added minus the number lost during a unit of time □ (Elzinga, 1981). Generally insect population had two distinctive characteristic. First many populations were extremely stable in size, generation after generation, second others could be extremely variable in size, with number rising rapidly in one or two generation to extremely high levels and then falling just as rapidly to very low numbers. □ (Ross, et.al. 1982). The two principal tactic of the r-strategy, nomadism and rapid population build up, clearly seen in plague locust, which are associated with areas all over the world □ (Matthews, 1984). K-strategic depend on many factor like availability of food and extent of predation □ ( Romoser, 1981), it’s happened in Morabine grasshopper in arid and semi-arid region of Australia. Basicly the r-/K- strategic based on assumption, that r- strategic have exponential population growth, stable age distributions, repeated colonization or fluctuations in population density and Kstrategy had logistic population growth environmental stable, population near equilibrium density, competition important Δ ( Stearns, 1977) Kind of insect which belongs to r- and K-strategy can be characterized below: r- strategy K-strategy Climate Uncertain variable, unpredictable Fairly consistent, reasonably certain. predictable, Life cycle Short, a few weeks Rapid development and growth rate Early reproduction and single reproduction Small body size Potentially many offspring Mortality catastrophic, usually Longer, usually several months Slower development and growth rate Delayed or extended reproduction period Large body size Fewer and large offspring Mortality more evenly distributed, density independent density dependent Population size Variable in time, non equilibrium, Fairly constant, equilibrium, at or near usually significantly below carrying carrying capacity, no need for capacity periodic recolonization recolonization Competition Variable commonly low Usually high. □ (Ross, et.al,1982 and Horn, 1976) References : - Ross, Herbert et.al. 1982. A textbook of entomology. John Wiley and Sons, Inc. USA. - Horn , David J.1976. Biology of insect. W.B. Saunders Company. Philadelphia, USA. - Matthews, E.G.1984. Insect ecology. University of Queensland Press. Australia - Elzinga, Richard J. 1978. Fundamentals of Entomology. Prentice Hall Inc. New Jersey. - Romoser, Wiliam S. 1981.Science of Entomology, second ed. Macmillan Publishing Co.Inc. New York - Stearn, Stephen C.1977. Annu.Rev.Ecol. Sys. 8:145-71 By Koko r-selection operates to maximize reproductive rate in harsh, unstable environments where population remain well below the carrying capacity of the environment are resources are not limiting except for time to reproduce successfully ( Peter, 1984).at the other extreme of the continuum, in predictable, stable environment populations will frequently reach the carrying capacity of the environment and selection will operate by improving adaptations for living in these crowded, competitive condition, where enemies such as predators and parasites will be very effective – k selection. The bare essentials of this theory of r- and k- selection therefore involve five characteristics that must be integrated there are: 1. Relatif stability of environmental conditions adn population size 2. Reproductive rate of species 3. Carrying capacity of the environment relative to mean population size. 4. The frequency of intraspecific and interspectif competition 5. The importance of enemy attack. In here can be explained the concept of r and k strategist and their development during the evolution of organism is certainly not new, although these particular terms to describe the process have been used for only a short period of time. There is presently a great deal of uncertainty as to how important these stratagems are in the functioning of organism within ecological communities. Dobzhansky, 1950 in Peter, 1974. He described that the differences between temperate or arctic environments and those of tropical regions. In tropical environment, organism are more constantly crowded and selection is influenced more by biotic factor. Competition then become a primary selective force because the populations are more constantly at or near the carrying capacity of the environment. MacArthur and Wison, (1967 in peter 1974) they said that the stage of the population there are two posible situations: (1) An environment in which a population is expanding, and (2) an environment in which a population is crowded. They assumed relatively stable condition in both cases, no great physical changes occurring to radically change the environment. Under this condition, organisms should evolve toward productivity and large families since there is no shortage of resourr-/KStrategy of insect Insects as another animals, has behavior to response the habitat or environment changes. Response to environment change and environmental changes produced the dynamical of insect it’s self. The dynamical population of insect known as r-/K- selection (another scientist called r-/K- strategist). r is symbol of rate of population growth and K is symbol population density. The number of individuals of species per unit area of space is referred to as population density, and growth of population is the number of individuals added minus the number lost during a unit of time □ (Elzinga, 1981). Generally insect population had two distinctive characteristic. First many populations were extremely stable in size, generation after generation, second others could be extremely variable in size, with number rising rapidly in one or two generation to extremely high levels and then falling just as rapidly to very low numbers. □ (Ross, et.al. 1982). The two principal tactic of the r-strategy, nomadism and rapid population build up, clearly seen in plague locust, which are associated with areas all over the world □ (Matthews, 1984). K-strategic depend on many factor like availability of food and extent of predation □ ( Romoser, 1981), it’s happened in Morabine grasshopper in arid and semi-arid region of Australia. Basicly the r-/K- strategic based on assumption, that r- strategic have exponential population growth, stable age distributions, repeated colonization or fluctuations in population density and Kstrategy had logistic population growth environmental stable, population near equilibrium density, competition important Δ ( Stearns, 1977) Kind of insect which belongs to r- and K-strategy can be characterized below: r- strategy K-strategy Climate Uncertain variable, unpredictable Fairly consistent, reasonably certain. predictable, Life cycle Short, a few weeks Rapid development and growth rate Early reproduction and single reproduction Small body size Potentially many offspring Mortality catastrophic, usually density independent Longer, usually several months Slower development and growth rate Delayed or extended reproduction period Large body size Fewer and large offspring Mortality more evenly distributed, density dependent Population size Variable in time, non equilibrium, Fairly constant, equilibrium, at or near usually significantly below carrying carrying capacity, no need for capacity periodic recolonization recolonization Competition Variable commonly low Usually high. □ (Ross, et.al,1982 and Horn, 1976) References : - Ross, Herbert et.al. 1982. A textbook of entomology. John Wiley and Sons, Inc. USA. - Horn , David J.1976. Biology of insect. W.B. Saunders Company. Philadelphia, USA. - Matthews, E.G.1984. Insect ecology. University of Queensland Press. Australia - Elzinga, Richard J. 1978. Fundamentals of Entomology. Prentice Hall Inc. New Jersey. - Romoser, Wiliam S. 1981.Science of Entomology, second ed. Macmillan Publishing Co.Inc. New York - Stearn, Stephen C.1977. Annu.Rev.Ecol. Sys. 8:145-71 This kind of evolution is called r- selection. On the other hand, under condition of crowding, selection should evolve toward feeding efficiency since resources are in short supply. The organism that can replace itself with the least waste of resources in the most fit in this environment, evolution here is called k selection. Pianka, (1970 in peter 1974) thoroughly treated the subject listed some of the correlates of both r and k selection. He propos that terestrial vertebrates and perennial plants are predominately k- selected. Whereas terrestrial invertebrates and annual plants are more r- selected. Gadgil and solbrig, (1972 in peter 1974) discussed r and k selection, objected to the assumption that increased or decreased birth brate by itself is evidence of increased r or k selection respectively. There are characteristics associated with both with both r and k strategists. Certain characteristic of r and k startegists is: R STRATEGISTS K STRATEGISTS Found most abudantly where density Found most abudantly where density independent factors predominate: dependent factors predominate: - temperate and arctic climate - tropical climate - edges population range - center population range - disturbed situation - undisturbed situation Have a high fecundity and short generation time Have a low fecundity and long generation time Usually reproduce early and rapidly, and died early. Usually reproduce early and rapidly, and died early. To be small in size To be large in size Dependent upon abudant food resources. Can exist with limited food resources highly competitive for resources. Poor competitors- opportunists good competitors- specialists May have greater tolerances to harsh environment conditions. May have restricted tolerances to harsh environment conditions. The concept of r and K strategies can act only as a mechanism for relative comparisons of biotic components of a community since there are no absolute criteria by which an organism can be designated as an r or K strategist. These comparisons may vary, depending on the species In www.springerlink.com r-strategist and k strategist can be classified according to their population growth curves ( MacArthur and Wilson, 1967 in Robinson). At one are the opportunists, the quantity breeders, that reproduce very large numbers of small-sized, biologically cheap, short-lived in individual. Each population explosion is followed by a population axtinction in which all active individuals normally perish and the species survives only in a dormant push as seeds, eggs, pupae, or spores. These quantity breeders is called r-strategist. Plant parasites, such as aphids are typical r-stategists. K-strategists depend on the survival ability of a complex individual. As the species subjected to r selection build up its numbers, and competitors and predators, the environment become crowded with species and the condition of r selection no longer prevail. These are the short of selectionpressures that operate in environment of the second kind, those that are predictably favourable and stable, the k environment ( k represents carrying capacity). Southwood, 1977a in Martin et all, elaborates on this set of evolutionary interactions with the environment. He associates the r-k spectrum with so called ‘’durational stability’’. Daly and howell, 1978 can describe the two principal tactict of the r-strategy, nomadism adn rapid population build up, on the locusts. The locust need condition favourable. These favourable condition are largely dependent on soil type. REFFERENCE Daly., howell.V. 1978. Introduction to Insect and Diversity. America. Martin, R. S., Mark, D.H, Allan, D.W. 1999. Ecology of insects concepts and applications. Department of zoology, university of Oxford. Oxford. Matthews, E. G., R.L. Kitching. 1976. Insect ecology. University of Queensland Press. Queensland. Peter, W.P. 1974. Evolutionary Strategies of Parasitic Insect. Department of Entomology University of Illinois Urbana, Illinois. Robinson, R. A. Host Management in Crop Pathosystem. University of California. Barkeley. www.springerlink.com