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Dispersal and persistence Disjunction – discontinuous range of a taxon in which at least two closely related populations are separated by a wide geographic distance 2 possible explanations 1. Dispersal 2. Geological events Dispersalists Darwin vs Extensionists Lyell Theory of Continental Drift Final evidence against extentionist arguments, but provided new theory critical to understanding species’ distributions Alfred Wegener Vicariance – geographical range of a taxa is split into parts by the formation of a barrier •Events such as plate tectonics, orogeny , climate change, sea level rise or glacial movement split populations Southern Beech Nothofagus Patagonia New Zealand • After vicariance events, evolutionary divergence can give rise to different (but related) species in the disjunctions Debate: Vicariance vs Dispersal What is the relative importance of long-distance dispersal events and vicariance (population-splitting events)? Eclecticist - dispersal and vicariance events are both important Definitions Ecological dispersal – movement of individuals away from an existing population or parent organism Reduce intra-specific competition But, habitat similarity decreases with distance Biogeographic dispersal – species shift their ranges by moving over long distances across large barriers (broader spatiotemporal scales) Infrequent but very important Mostly historic examples Why disperse? • Reduce intraspecific competition • Find more suitable habitats Successful range expansion 1. Travel to new area 2. Tolerate conditions of new habitat 3. Reproduce in new habitat Mechanisms of dispersal Active: movement of an organism from one location to another by its own means Passive: movement of an organism from one place to another by means of a stronger force, such as water flow, wind or another organism Monarch Butterfly Golden Plover Active dispersal – Air Some species have extensive migratory routes and thus show great potential for dispersal Ruby-throated hummingbird Active dispersal - water Himalayan Balsam Squirting cucumber Active dispersal - plants Passive dispersal - water Passive dispersal - water Passive dispersal – floating islands Passive dispersal – wind Diaspore – plant dispersal unit (consists of seed plus additional tissues adapted for dispersal) Pappus Wind dispersal of large propagules Wings Exozoochory – seeds dispersed on outside of animal Endozoochory – seeds dispersed on inside of animal Types of dispersal events 1. Jump dispersal 2. Diffusion 3. Secular migration Jump dispersal Long distance dispersal accomplished during a relatively short period – Occurs infrequently, large effect – Some species are better at jump dispersal than others (water, wind, wings) e.g. Hawaii and Galapagos Consequences: 1. 2. Explains wide and often discontinuous distributions of many taxa Accounts for similarities/ differences among biotas inhabiting similar environment in different geographic areas (some species good at distance dispersal, but chance is still important) Human-aided jump dispersal increasing Diffusion Slow dispersal of individuals spreading out from the margins of the species’ range (accomplished over generations) Often follows jump dispersal Cattle egret Jump dispersal + diffusion Africanized honey bees Brought from Africa Escaped from Sao Paolo Dispersed ~ 100 mi/yr Secular migration Very slow dispersal (e.g. hundreds of generations) that commonly involves evolutionary changes in the dispersing populations Secular migration - camels Guanaco Vicuña Barriers and Dispersal Routes Barriers – abiotic or biotic feature that restricts movement of genes or individuals from one place to another • Species-specific • Generally, organisms that inhabit fluctuating environments are more tolerant of extreme/unusual conditions than species in stable habitats Janzen (1967) Types of Barriers Physiological Ecological Behavioral Physiological Barriers Physical environments outside the range an organism normally encounters so it cannot survive and disperse across the barrier Common physiological barriers – Land-water – Salinity – Temperature Ecological Barriers Dispersing organisms must be able to withstand ecological hazards – Predation – Competition Behavioral Barriers Ability of some organisms to select suitable habitat can limit ability to disperse across barriers – Example – bird species capable of flying long distances will not cross open areas Dispersal Routes Corridors Filters Sweepstake routes Increasing resistance to biotic exchange Corridor – dispersal route that permits movement of many related species from one region to another – Similar environment to that of the two regions – Balanced taxonomic composition Beringia - Pleistocene Proposed I-70 corridor (unfunded) Banff Corridor Filters - selectively impede the movement of certain taxa while allowing others to disperse freely (biotic or abiotic) – Colonist-biased subset – Often form transition zones Filters Two-way filter for reptiles between Australia and Asia Sweepstake Routes – severe barrier that permits rare dispersal events (jump dispersal) • Rare on ecological timescale, likely and important on geological timescale • Can be followed by dispersal and adaptive radiation 1995, Hurricane Luis Vegetation mats Green iguanas (Iguana iguana) sweepstakes route from Guadeloupe to Anguilla Corridors Favorable habitat connecting larger ones Filters Blocks or slows passage of some organisms Sweepstake Routes Severe barrier that permits rare dispersals Successful range expansion 1. Travel to new area 2. Tolerate conditions of new habitat 3. Reproduce in new habitat Once dispersed to new habitat, species may select appropriate habitat Mobile organisms – search for favorable habitats Immobile organisms – some do not settle or germinate unless perceive certain sensory cues Which organisms often have good survival in new ecosystems? From highly fluctuating/unpredictable environment From large, ecologically-diverse habitats Successful range expansion 1. Travel to new area 2. Tolerate conditions of new habitat 3. Reproduce in new habitat Reproductive traits of “good colonizers” Asexual reproduction (need only one) Vegetative reproduction (ramets, bulbs) Apomictic (seeds from unfertilized ovules) Parthenogenesis (embryos from unfertilized eggs) Sexual reproduction Hermaphroditic and self-compatible High reproductive rate, short gestation