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Notes and Examples for LIFE HISTORY - the pattern of reproduction, growth, maturation and dispersal that generates an organism’s FITNESS The problem is one of ALLOCATION: How does an organism allocate limited resources? (It can’t do everything.) FITNESS = REPRODUCTIVE OUTPUT X When/how often to reproduce When to cease reproduction Energy required for displays - behavioral - morphological How many offspring to produce: - per brood - per lifetime Invest in parental care, or not PROBABILITY OF SURVIVAL How long to continue growth: - short time (end up small) - long time (end up large) Predator defences Disease resistance Special skills, e.g. competitive ability Dispersal: - when? - how often? - how far? WHAT ARE THE ADVANTAGES OF BEING LARGE OR SMALL? LARGE ORGANISMS • • Slow metabolic rate; low energy demand per unit biomass (efficient) Need less food SMALL ORGANISMS • High metabolic rate; high energy demand per unit biomass (less efficient) • Need more food • Relatively well buffered against abiotic fluctuations • Usually more sensitive • Mass, aggressiveness and weaponry are possible options; advantage in a competitive world • These options not so available • Long life with multiple episodes of reproduction (“iteroparity”) • • Low rmax values • Short life, few episodes of reproduction (in the extreme, one episode – “semelparity”) High rmax values So overall: IT PAYS TO BE SMALL WHEN TIMES ARE GOOD - LOTS OF FOOD, LOW FLUCTUATIONS ETC.....BUT....WHEN TIMES ARE BAD, LARGE ORGANISMS HAVE THE ADVANTAGE. Therefore: (A). Under conditions of INTENSE COMPETITION for resources, individuals that divert energy to competitive abilities (growth, defense, etc.) will leave more successful offspring than those merely continuing to reproduce. • i.e. K-selection characteristics have selective advantage under intense competition. (B). By necessity, such organisms must have LOW rmax values. • i.e. they have higher ra values (not rmax) under competitive environments than "reproducers" Note: a K-strategist will leave fewer offspring under intense competition than under low competition, but under intense competition will leave more than an r-strategist. (C). Under conditions of LOW COMPETITION, individuals that devote resources to reproduction as soon as possible will leave more successful offspring than those who devote energy to growth etc. - best strategy is to keep 'G' (generation time) small. • i.e. r-selection characteristics have selective advantage under low competition. CHARACTERISTICS OF SPECIES WITH C, S, & R STRATEGIES* The expected life-history traits will be C-, S-, or R-selected types…: When frequency/ intensity of disturbance is: When harshness/stress is: LOW HIGH LOW C S HIGH R - C - SELECTION: COMPETITORS - selected for high competitive ability which depends upon plant characteristics which maximize the capture of resources in productive, relatively undisturbed environments. S - SELECTION: STRESS-TOLERATORS - has brought about reductions in both vegetative and reproductive vigour, adaptations which allow endurance of continuously unproductive environments e.g. arctic, alpine or arid environments. R - SELECTION: RUDERALS - associated with short life span and high seed production in highly disturbed but potentially productive environments e.g. arable situations. DEFINITIONS: Stress (=Harshness) – any factor which slows or limits the ability to accumulate biomass Disturbance – any factor which removes units of biomass and/or whole individuals *GRIME, J.P. (1977). Evidence for the existence of three primary strategies and its relevance to ecological and evolutionary theory. American Naturalist 111:1169-1194. Genet – physically separate unit of plant biomass with a genotype different from that of its neighbouring genets; may be composed of a single ramet, or many Ramet (=“Tiller”) – an attached or separate component of a genet; all ramets of a given genet are identical in genotype (but likely variable in phenotype, especially size)