Lecture 11 – LIFE HISTORY STRATEGY Life History Strategy A suite of traits that improve an individual's chances of surviving and reproducing in a particular environment COST/BENEFIT ANALYSIS 1. Energy committed to one function is not available for others. Growth Respiration Reproduction Predator Defense Growth Respiration Reproduction Predator Defense COST/BENEFIT ANALYSIS 1. Energy committed to one function is not available for others. 2. Every evolutionary ‘choice’ has costs and benefits Settles on wave exposed shore Costs Benefits -energy in holding on -less exposure to predators -restricted movement for food gathering -more food delivery -size restriction -less energy directed to growth COST/BENEFIT ANALYSIS 1. Energy commited to one function is not available for others. 2. Every evolutionary ‘choice’ has costs and benefits 3. Every organism represents a compromise of a number of evolutionary ‘choices’ Periclimenes yucatanicus Colour - bright & obvious to predators - disruptive Hides in anemone – restricted foraging but has protection Exoskeleton – restricts movement but offers protection Macarthur and Wilson (‘70’s) - Theory of Island Biogeography r strategists Opportunistic -rapidly colonizing, fast growth K strategists R.H. Macarthur E.O. Wilson Equilibrium -slower colonizing, slow growth Attributes of r and K - selected organisms r - selected Climate Mortality Population Size K - selected Variable, unpredictible Consistent, predictible Density independent, uncertain adult survival Density dependent, uncertain juvenile survival Variable, below K Consistent More species, no recolonization Community Characteristics Few species, recolonization Lifespan Short Individual Attributes Rapid development, early reproduction, high rate of increase, small body size Slow development, late reproduction, low rate of increase, larger body size Allocation of Energy Greater % to reproduction Greater % to growth and maintenance Long Attributes of r and K - selected organisms r - selected K - selected Climate Variable, unpredictible Consistent, predictible Mortality Density independent, uncertain adult survival Density dependent, uncertain juvenile survival Population Size Variable, below K Consistent Community Characteristics Few species, recolonization More species, no recolonization Lifespan Short Long Individual Attributes Rapid development, early reproduction, high rate of increase, small body size Slow development, late reproduction, low rate of increase, larger body size Allocation of Energy Greater % to reproduction Greater % to growth and maintenance Productivity Attributes of r and K - selected organisms r - selected K - selected Climate Variable, unpredictible Consistent, predictible Mortality Density independent, uncertain adult survival Density dependent, uncertain juvenile survival Population Size Variable, below K Consistent Community Characteristics Few species, recolonization More species, no recolonization Lifespan Short Long Individual Attributes Rapid development, early reproduction, high rate of increase, small body size Slow development, late reproduction, low rate of increase, larger body size Allocation of Energy Greater % to reproduction Greater % to growth and maintenance Productivity Efficiency Look at our original energy allocation graphs Growth Growth Respiration Respiration Reproduction Reproduction Predator Defense More r - like Predator Defense More K - like r - K continuum r-strategist Frequent disturbance K-strategist Less frequent disturbance Aclonal Life History recruitment Sexual reproduction Death Clonal Life History Sexual reproduction recruitment Fission Fission Fission Death Life history pattern -- a genetically inherited pattern of resource allocation to that optimizes the passing of genes to the next generation EGG AND SPERM PRODUCTION FERTILIZATION PATTERNS TYPES OF DEVELOPMENT LARVAL DISPERSAL STRATEGIES SETTLEMENT PATTERNS Fertilization Patterns 1. External (no copulation) Fertilization Patterns 1. External (no copulation) Problems Gamete output Synchrony Gamete dispersal Gamete dilution Fertilization Patterns 1. External (no copulation) Problems Gamete output Synchrony 100 % success Gamete dispersal Gamete dilution 0.5 m Distance between individuals Fertilization Patterns 1. External (no copulation) Allee Effect Reproductive rate Number or Density Fertilization Patterns 2. Internal (no copulation) Fertilization Patterns 2. Internal (copulation) Fertilization pattern Costs External Fertilization -large gamete output Benefits -takes advantage of medium -need for synchrony -unpredictable dispersion -gamete dilution Internal Fertilization (no copulation) -large gamete output -takes advantage of medium -unpredictable dispersion -gamete dilution Internal Fertilization (copulation) -energy in mate searching -less gamete wastage -energy in courtship -greater efficiency in reproduction 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Overview – Life History Patterns 1) Fertilization patterns ✓ 2) Development patterns 3) Dispersal patterns 4) Settlement patterns 4) Settlement patterns Developmental Patterns -Kinds of eggs Isolecithal • •• • •• • • •• • • •• •• • •• • • Telolecithal ••••••••••• •• • • • • • •••••••••••••••••• ••• • •• • • yolk 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Developmental Patterns -Kinds of eggs 4) Settlement patterns Isolecithal • •• • •• • • •• • • •• •• • •• • • Cleavage through entire egg Telolecithal ••••••••••• •• • • • • • •••••••••••••••••• ••• • •• • • Cleavage not through entire egg 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Developmental Patterns -Kinds of eggs 4) Settlement patterns Holoblastic Isolecithal • •• • •• • • •• • • •• •• • •• • • Cleavage through entire egg Telolecithal ••••••••••• •• • • • • • •••••••••••••••••• ••• • •• • • • • •• • •• • • ••• ••••• ••• •••• • ••• •• ••• • • •• • •• Meroblastic Cleavage not through entire egg •••• •• ••••••• ••••••••• •• •• • • • • •• Developmental Patterns -Kinds of eggs Isolecithal - Holoblastic 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns Telolecithal - Meroblastic 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Developmental Patterns -Kinds of eggs Isolecithal • •• • •• • • •• • • •• •• • •• • • Telolecithal ••••••••••• •• • • • • • •••••••••••••••••• ••• • •• • • 4) Settlement patterns Holoblastic •• •• ••• • ••• • • • ••••• • •••• • ••• •• ••• • • •• • •• Planktotrophic larvae Meroblastic •••• •• ••••••• ••••••••• •• •• • • • • •• Lecithotrophic larvae 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Kinds of Development Patterns 4) Settlement patterns Free spawning Weakly isolecithal egg Indirect Maturation Free spawning Planktotrophic, free-swimming larvae Settlement and metamorphosis Indirect Maturation Juvenile Mating Brooding of embryos Direct Maturation Settlement and metamorphosis Juvenile Mating Strongly telolecithal egg Planktotrophic, free-swimming larvae Strongly/moderately telolecithal egg Hatch as juveniles Moderately telolecithal egg Maturation Juvenile Brooding of embryos Mixed Hatch as freeswimming larvae Settlement and metamorphosis 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Larval Settlement – Large Scale 4) Settlement patterns 0.2 % recovery 0.2 % recovery 0.18 % recovery Scheltema ‘71 Larval Settlement – Large Scale 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns Scheltema ‘71 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Larval Settlement – Large Scale 4) Settlement patterns 1012 1010 Frequency of dispersal 108 106 104 102 102 104 106 108 1010 1012 Population size Scheltema ‘71 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Larval Settlement – Large Scale 4) Settlement patterns Volume of water (m3) Total number of larvae North Atlantic Drift 2.45 x 105 3.31 x 1011 North Equatorial Current 3.43 x 105 4.63 x 1011 South Equatorial Current 2.26 x 105 3.05 x 1011 Equatorial Undercurrent 0.98 x 105 1.32 x 1011 Scheltema ‘71 Larval Settlement Large Scale Point Conception 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Larval Settlement Large Scale 4) Settlement patterns Point Conception Current Patterns Larval Settlement Large Scale 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns Point Conception Current Patterns Biogeographical Zones Embayment Flushing (Gaines & Bertness, ‘92) 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns Wind & Tidal Currents 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns Associated with internal waves 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Settlement - Behaviour of Larvae 4) Settlement patterns Metamorphosis to postlarva Drifts in plankton (days/weeks) Lobster hatches -photopositive- Preference switches to photonegative Lobster settles and undergoes metamorphosis 3. Benthic/Surface Phenomena 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns Pawlik et al, ‘91. Science 251:421 - settlement of reef-building worm Current Speed -low -medium -high (>15 cm/sec) Phragmatopoma 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 3. Benthic/Surface Phenomena 4) Settlement patterns Pawlik et al, ‘91. Science 251:421 No ‘sampling’ # settling Current Erosion 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Early models 4) Settlement patterns Larvae in water column Settle randomly Differential mortality Adult distribution More recent view Larvae in water column Passive deposition Testing substrate Not settle Settle Adult distribution 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns Passive Deposition -due to 1) sedimentation rate 2) swimming behaviour 3) bottom flow patterns 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns Bottom flow patterns Laminar flow 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns Bottom flow patterns Change in angular velocity x xx x x Experiment on settling and flow Botman et al 1998 Capitella rich muds Mercenaria coarse sand Experiment on settling and flow Flow Glass beads Mud Experiment on settling and flow Still water Mud Capitella Mercenaria Flow Beads yes no Conclusion Mud beads no yes no yes No preference No preference - choice is species specific - Mercenaria couldn’t ‘sample’ 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns General Model 4) Settlement patterns Free swimming larva Alternating photonegative & photopositive behaviour Passive deposition and contact with surface Surface texture Chemical cues Attachment Contact with conspecifics 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns Larval Settlement 1. Conspecifics 2. Microbial films 3. Prey species 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Larval Settlement 4) Settlement patterns 1. Conspecifics Free fatty acids Larval settlement Phragmatopoma 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Larval Settlement 4) Settlement patterns 2. Bacterial films Polysaccharides, Glycoproteins Increased searching or metamorphosis Larval Settlement 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns 4) Settlement patterns 3. Prey species - herbivores Settlement induced by GABA 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Larval Settlement 4) Settlement patterns 3. Prey species - herbivores Gigartina Abalone larva In field Porphyra -not found on either -not induced to settle on whole fronds 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Larval Settlement 4) Settlement patterns 3. Prey species - carnivores Onchidoris bilamellata Induces settlement 1) Fertilization patterns 2) Development patterns 3) Dispersal patterns Larval Settlement 4) Settlement patterns 3. Prey species - carnivores Phestilla Induces settlement < 500 Da protein (< 10 mM)