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Biology 450: Fish Week Lecture 11: Ontogenetic shifts and dispersal Scott Heppell Department of Fisheries and Wildlife Oregon State University 042 Nash Hall 737-1086, [email protected] Ontogenetic shifts Fishery Ontogeny: Biology The entireFall development 2006 of an organism, from fertilization through maturity The salmon life cycle http://www.streamnet.org More than salmon … More than benthic species… © Richard Herrmann, Poway, California, http://www.richardherrmann.com Types of ontogenetic shifts • • • • Habitat/location/migration Reproductive strategy Diet Behavior Ontogenetic shifts: change in lifestyle associated with growth and development • • • Growth rates in habitat x slow, so make a shift to habitat y Maximize growth, minimize mortality risk: m/g Major shifts common in species with very large changes in body size – – – – Larvae Juveniles – Increase in depth with body size Adults Spawning adults Why switch habitats? (Werner and Gilliam 1984) Growth Rate (mm/day) Optimal switching point Habitat B Habitat A Body size Mortality risk/growth rate (m/g) Want to minimize mortality (m) AND maximize growth (g) Habitat A Habitat B Body size Loggerhead sea turtle growth curve (mark recapture estimates or width of growth rings on humerus) Length Adult Subadult Pelagic Benthic juvenile AGE Migration Definition: a move from one place to another 3 Types • Local and seasonal movements • Dispersals • True migrations Larvae: pelagic Small juveniles: estuaries, kelp forests Adults: deeper reefs Larger juveniles: nearshore reefs 1. Local and Seasonal Movements Involves • short or long distance • seasonal to semidiurnal time periods • horizontal or vertical movements Migratory records • Mather (1962) tagged two bluefin on June 1 and June 10 in waters off the Bahamas. • On September 28 and October 6, respectively, these two fish were recovered near Bergen, Norway. 4000 nm 2. Dispersal Includes • Passive or active movement away from the breeding habitat over a wide area (e.g., larval transport, juvenile dispersal) • Ontogenetic shifts between habitats 3. ‘True’ Migration Involves • Directed movement between widely separated and well-defined areas, often at set times during the year (e.g., feeding to breeding areas) • Often ignores local conditions Why migrate? • Breeding – Aggregations for mating/spawning – Optimal birthing/nesting areas Why migrate? • Breeding – Aggregations for mating/spawning – Optimal birthing/nesting areas • Feeding – Higher latitudes = more food – Food patches often ephemeral http://earthobservatory.nasa.com Migration to Spawning Sites & Passive or Active Larval Migration • Improve breeding success • Reduce impact of environmental fluctuations • Larval Dispersal • Cushing’s (1957) match-mismatch hypothesis Evolutionary Perspective Migration involves specialized behaviors that have arisen through natural selection How do we think about this? Trade Offs: Costs versus Benefits Benefits • Increased feeding opportunities • Avoidance of adverse environmental conditions • Improved current reproductive success Costs • Energetic cost of migration itself • Increased predation risks • Energetic & development costs of any specific migratory adaptation • Potential reproductive costs due to decreased lifetime reproductive effort Fitness Summation of: Probability of survival to age x lx Multiplied by Fecundity and breeding success mx Migration favored if: S lxmx migrators > S lxmx residents Migration - Maturation Decision • Given that migration is costly and potentially dangerous, age or size at maturity may be delayed • Some species (some rockfish, sturgeon, others?) appear to go through a “false maturation” in which females physiologically prepare for reproduction but resorb their eggs instead of completing maturation Migratory Patterns • Oceanodromous • Potamodromous • Diadromous Diadromy Gross ‘87 McDowall ‘87 Gross et al. ‘88 Migration pathways of large pelagics • Can follow physical features, such as continental margins or major currents • However, “corridor” concept probably too simplified – Individual variability – Variation in response to ocean conditions • Behavior depends on species, time of year, sex, other factors – Coastal, pelagic – Direct, stop-and-feed • Moving oases of food resources require behavioral plasticity Movements of Leatherback Sea Turtles in the Atlantic Ocean as Determined Using Satellite Transmitters Why might understanding migration pathways be important from a management perspective? Hector Barrios-Garrido NOAA Lutcavage et al. ‘99 Legend: 1998, 1999, Early Jettison March/April May June July Oceanodromous (European Sea Bass, Dicentrarchus labrax) Overwintering Spawning Aggregation spawning Why aggregate? • • • • Access to mates Multiple fertilizations Environmental conditions Reduced offspring mortality Management • Ontogenetic shifts in habitat type dictate that habitat protections be in place for all life stages • Highly migratory species do not care about EEZ lines international cooperation and research is required • Bycatch in high seas fisheries presents a management and monitoring problem – need for integration among groups – Impact may be large but diffuse and therefore difficult to measure Options? • Fishing and bycatch quotas • Gear modification • Closures – Spatial, temporal – Requires consideration of entire life cycle, migration rates and habitat use – Requires careful consideration of redistribution of effort and markets Modeling movement for reserve planning • Understanding of: – – – – Movement patterns Reproductive strategy Location of impacts Suite of management options Types of movement Habitat A “leaking” = stochastic movement Habitat B “reserve” Types of movement Habitat A “spill-over” = densitydependent leaking Habitat B “reserve” Types of movement Habitat A “seeding” = larval transport Habitat B “reserve” Types of movement Habitat A Habitat B “reserve” “ontogenetic shift” = deterministic movement between habitats due to life stage transition RESERVE NON-RESERVE Offspring production Survival and growth Spillover or leaking Life stage with increased survival Newborn juvenile Adult Newborn juvenile Adult RESERVE NON-RESERVE Offspring production Survival and growth Spillover or leaking Egg/larvae juvenile Ontogenetic shift Seeding from reserve Spawning Adult Non-spawn adult Spawning adult RESERVE NON-RESERVE Offspring production Survival and growth Spillover or leaking Egg/larvae juvenile juvenile Ontogenetic shift Adult RESERVE NON-RESERVE Offspring production Survival and growth Spillover or leaking Egg/larvae Ontogenetic shift Seeding from reserve juvenile Adult juvenile Adult RESERVE NON-RESERVE Offspring production Survival and growth Spillover or leaking Ontogenetic shifts Seeding from reserve Egg/larvae juvenile Adult Spawning adult Egg/larvae juvenile Adult Spawning adult Generic model Conclusions • Ontogenetic shifts happen as individuals try to maximize µ/g • Shifts happen in habitat selection, reproduction, diet, and behavior • Incorporating an understanding of these shifts crucial to effective management and conservation • Plasticity in ontogeny is important but not always well understood – Manage for variance, not just the mean