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5 Reproduction, Dispersal, and Migration
Notes for Marine Biology: Function, Biodiversity, Ecology By Jeffrey S. Levinton ©Jeffrey S. Levinton 2001
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Sex and Reproduction IS SEX NECESSARY?
WE MUST SEPARATE SEX AND REPRODUCTION? SPECIES CAN REPRODUCE WITHOUT SEX (CLONAL GROWTH INVOLVING FISSION OR BUDDING OF INDIVIDUALS)
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Sex and Reproduction Non-sexual reproduction:
Descendants are genetically identical - clone Colonial species produce a set of individuals that are genetically identical, known as a module; each module may have arisen from a sexually formed zygote
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Cost of Sex FEMALE gives up half her possible genes in progeny
Sex involves expenditure of energy and time to find mates, combat among males
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Benefits of Sex? genetic diversity - sex increases combinations of genes - resistance against disease Alternative to sex: clones, must wait for mutations to occur Sex - recombination produces variable gene combinations, meiosis enhances crossing over of chromosomes: new gene combinations and intragenic variants
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Sexual Selection selection for extreme forms that breed more successfully - major claw of fiddler crabs, deer antlers, colors of male birds Can involve selection for display coloration, enhanced combat structures Female choice often involved; selection for fit males (good genes hypothesis)
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Sexual Selection The major claw of fiddler crabs is employed
for display to attract females and for combat with other males
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Types of Sexuality Separate sexes -gonochoristic
Hermaphroditism -individual can have male or female function
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Hermaphroditism Simultaneous Sequential
Protandrous - first male,then female Protogynous - first female then male
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Sequential Hermaphroditism
Protandry - size advantage model Eggs costly in terms of resources, so more offspring produced when individual functions as female when large Male function does not produce great increases in offspring when it gets larger Therefore, there is a threshold size when female function begets more offspring.Smaller individuals do better as males.
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Male at advantage Female at advantage
Number of offspring produced Male Body size The size advantage model for Protrandry
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Protogyny Male function must result in more offspring when male is older and larger Important when aggression is important in mating success, e.g., some fishes where males fight to maintain group of female mates
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Male polymorphism Males may occur as aggressive fighting morphs, or less aggressive morphs Found in a number of groups, e.g., some fishes and some amphipod or isopod crustaceans Determination of morphs can be environmental, genetic Less aggressive morphs can obtain mates by “sneaky” tactics, which are often successful
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Factors in Reproductive Success
Percent investment in reproduction - reproductive effort Age of first reproduction (generation time) Predictability of reproductive success Juvenile versus adult mortality rate
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Life History Theory Tactics that maximize population growth
Evolutionary “tactics”:Variation in reproductive effort, age of reproduction, whether to reproduce more than once Presume that earlier investment in reproduction reduces resources available to invest in later growth and survival
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Examples of Life History Tactics
Strong variability in success of reproduction:reproduce more than once High adult mortality: earlier age of first reproduction,perhaps reproduce only once Low adult mortality: later age of first reproduction, reproduce more than once
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Example: Selection in a Fishery
Shrimp Pandalus jordani,protandrous Danish, Swedish catch (Skagerak) – stable, increased slowly – catch tripled (2000 6300 ton/y)
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Pandalus jordani fishery
Changes in Body Size Changes in Size of Change from Male to Female
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Sex - factors in fertilization
Planktonic sperm: (and eggs in many cases). Problem of timing, specificity. Direct sperm transfer: (spermatophores, copulation). Problem of finding mates (e.g., barnacles, timing of reproductive cycle)
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Planktonic sperm and eggs
Specialized binding/fertilization proteins in sperm and receptors in eggs (bindin in sea urchin sperm, lysin in abalone sperm) Sperm attractors in eggs Binding proteins are species-specific, proteins with high rates of evolution
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Gamete matching important in plankton
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Timing of sperm and egg release
Epidemic spawning - known in mussels, stimulus of one spawner causes other individuals to shed gametes Mass spawning - known in coral species, many species spawn on single nights Timing of spawning (also production of spores by seaweeds) at times of quiet water (slack high or low tide) to maximize fertilization rates
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Movement of Marine Organisms
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Dispersal versus migration
DISPERSAL: UNDIRECTED MIGRATION: DIRECTED, RETURN SPECIFIC
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Migration scheme Adult Stock Spawning Area Nursery/Juvenile
Feeding Area
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Migration Types ANADROMOUS - fish live as adults in salt water, spawn in fresh water (shad, striped bass), more common in higher latitudes CATADROMOUS - fish live as adults in fresh water, spawn in salt water (eel) more common in lower latitudes FULLY OCEANIC - herring, green turtle
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Migration
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Norway Migration of the herring in the North Sea Adult feeding area Spawning areas
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N. America Europe Africa
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Larval Dispersal
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Dispersal Types in Benthic Species
PLANKTOTROPHIC DISPERSAL - female produces many ( ) small eggs, larvae feed on plankton, long dispersal time (weeks), some are very long distance (teleplanic) larvae - cross oceans LECITHOTROPHIC LARVAE - female produces fewer eggs ( ), larger, larvae live on yolk, short dispersal time (hrs-days usually) DIRECT RELEASE - female lays eggs, or broods young, juveniles released and crawl away
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Lecithotrophic larva: tadpole larva of the
colonial ascidian Botryllus schlosseri Pluteus larva of an urchin Planktotrophic larva of snail Cymatium parthenopetum
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Loss to offshore waters
Wind-driven recruitment onshore Internal waves, tidal bores Self-seeding eddies Longshore drift Shore Population
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Some helping hands in dispersal
Winds that wash larvae to shore Internal waves - bring material and larvae to shore Eddies that concentrate larvae in spots Behavior - in estuaries can allow retention (rise on the flood tide, descend on the ebb tide)
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Estuarine larval adaptations - retention
Larvae rise on the flooding tide, sink to bottom on the ebbing tide: results in retention of larvae within estuary
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Estuarine larval adaptations - movement of larvae to coastal waters, return of later stage larvae
Blue crab, Callinectes sapidus
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Effect of local eddies on larval retention in a patch reef on the Great Barrier Reef, Australia
Planula larva Recruitment of juvenile corals Distance from reef perimeter Newly settled coral
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Why disperse? High probability of local extinction; best to export juveniles Spread your young (siblings) over a variety of habitats; evens out the probability of mortality Maybe it has nothing to do with dispersal at all; just a feeding ground in the plankton for larvae
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Settling problems of planktonic larvae
Presettling problems: Starvation Predation in plankton Loss to inappropriate habitats
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Example of Effect of Starvation:
Phytoplankton variation and barnacle larval success Abundant diatoms 1950 Normal phytoplankton 1000 500 Cyprids settling Early Larval stages Later Larval Stages Number of larvae 1951 Failure of phytoplankton Diatom failure March April Semibalanus balanoides settlement in a Scottish Sea Loch
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Postsettling problems
Energetic cost of metamorphosis Predation Crowding --> mortality Initial 6 months 12 months Expectation of life (months) 18 months 2 Interindividual contacts per cm Expectation of life of Semibalanus balanoides as function of crowding
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Stages in the selection of substratum by planktonic larvae
Free-swimming larva Selection Behavior Alternating Photo+ and Photo- stages Block in behavior, contact with inappropriate surface Releasor Random contact With a surface Selection behavior- crawling and test surfaces Releasor Releasor Releasor Block in behavior, e.g., contact with crowded surfaces Contact w. Substance on Sfc. Of another species Contact w. pits and grooves Contact w. adults of same sp. Selection behavior- frequent turning and flexing ATTACHMENT Stages in the selection of substratum by planktonic larvae
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The End
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