Speciation: A Diagram Book By Adam Rothstein For Bordertown Speciation: A Diagram Book Welcome to the Speciation Diagram Book! Speciation is the evolutionary process by which species derive. This crucial theory to the spread of new species is a little tricky, however. This handbook has been designed to illustrate the concepts of speciation, by animating the interactions of animal populations in low-tech, flipbook style diagrams. All theories of speciation, in that they represent ' cladogenesis' or the 'cleaving' of species into separate lineages, rather than ' anagenesis', or evolution within lineages, occur in relation to a topological boundary. The diagrams in this book are tripartite: there is a left portion, a right portion, and a center portion. The center denotes a ' species border'. Whether geographic, topological, or behaviorially reified, this border is the splitting point. It is the point at which individuals are forced to make their mating choices, and the point at which the gene pool of species has a change to mimic this boundary line. Watch this space. New borders are evolving all around us, and life evolves along with it. 1 First, let's talk about... Genetic Drift Although the exact role played by genetic drift in speciation is uncertain, it is a potential explanation for how cladogenesis speciation occurs rapidly, whereas anagenesis is a much slower process. Natural selection is a well-known means of evolution. Over time, improvements in individuals' genetic fitness increase the prevelance of certain traits, as those individuals more successful at breeding contribute their genetic makeup to the total population. Genetic drift, on the other hand, can potentially effect large changes in the gene pool much quicker than otherwise would happen naturally, due to random sampling. The gene pool is only defined by those individuals in the population that interbreed. A rare characteristic can become more common quickly, if random sampling gives that characteristic a larger than average chance at being represented within the breeding population. 1 First, let's talk about... Genetic Drift Although the exact role played by genetic drift in speciation is uncertain, it is a potential explanation for how cladogenesis speciation occurs rapidly, whereas anagenesis is a much slower process. Natural selection is a well-known means of evolution. Over time, improvements in individuals' genetic fitness increase the prevelance of certain traits, as those individuals more successful at breeding contribute their genetic makeup to the total population. Genetic drift, on the other hand, can potentially effect large changes in the gene pool much quicker than otherwise would happen naturally, due to random sampling. The gene pool is only defined by those individuals in the population that interbreed. A rare characteristic can become more common quickly, if random sampling gives that characteristic a larger than average chance at being represented within the breeding population. A sudden die-off, a splitting of one herd or flock into two, or any other separation that results into one large breeding group dividng into multiple smaller breeding groups can result in an individual or individuals with a particular trait formerly in the minority suddenly becoming much more representative throughout the population. What was previously rare in terms of the whole, is now more common. Individuals with that rare trait might have previously fought a hard fight to breed and continue adding the trait to that population's gene pool. But now, with competitors reduced or new territory available, they might find it easier to breed, and so their minority increases in number relative to the whole. A sudden die-off, a splitting of one herd or flock into two, or any other separation that results into one large breeding group dividng into multiple smaller breeding groups can result in an individual or individuals with a particular trait formerly in the minority suddenly becoming much more representative throughout the population. What was previously rare in terms of the whole, is now more common. Individuals with that rare trait might have previously fought a hard fight to breed and continue adding the trait to that population's gene pool. But now, with competitors reduced or new territory available, they might find it easier to breed, and so their minority increases in number relative to the whole. Perhaps, once there is a more sizeable number of individuals expressing that trait, the trait's improvements to the genetic fitness of the individuals begin to take hold. Maybe animals with improved speed hunt better together, whereas a single faster individual only made better escapes from predators. What genetic drift started, natural selection continues, until the minority trait is found in a majority of the population. Additionally, competition with the individuals bearing the new trait might make life harder for those without it, who survived fine previously, but now struggle against the up-and-comers. Perhaps the trait is a new marking pattern, or significantly better survival behavior that leaves the other individuals more exposed to predators. In a sense, these "obsolete" individuals are at the mercy of the same curve from which they previously reaped the reward at the expense of others. Evolution is about competition, and competition itself is a state that begets multiples in benefits for winners and hazards for losers. Perhaps, once there is a more sizeable number of individuals expressing that trait, the trait's improvements to the genetic fitness of the individuals begin to take hold. Maybe animals with improved speed hunt better together, whereas a single faster individual only made better escapes from predators. What genetic drift started, natural selection continues, until the minority trait is found in a majority of the population. Additionally, competition with the individuals bearing the new trait might make life harder for those without it, who survived fine previously, but now struggle against the up-and-comers. Perhaps the trait is a new marking pattern, or significantly better survival behavior that leaves the other individuals more exposed to predators. In a sense, these "obsolete" individuals are at the mercy of the same curve from which they previously reaped the reward at the expense of others. Evolution is about competition, and competition itself is a state that begets multiples in benefits for winners and hazards for losers. Individuals of a species are always competing, not just against other species for habitat and resources, but against the members of their own species, even as they work together to breed and continue their population. Eventually, the prior trait may be wiped out of existence. The random sampling of genetic drift promotes traits by simply increasing their relative odds. After that point, natural selection runs its course, but with a different beginning point than might otherwise have been. Both genetic drift and natural selection play into the evolution of species, but as we will see, geographic speciation creates conditions under which genetic drift is likely to occur, accelerating the rise of new mutations to the top of the expressionary gene pool heap. Individuals of a species are always competing, not just against other species for habitat and resources, but against the members of their own species, even as they work together to breed and continue their population. Theories ofSpeciation.... Allopatric Speciation Allopatric speciation (from the ancient Greek allos, "other" + Greek patra, "fatherland") occurs when a population is split into two geographically isolated populations, such that they are able to interbreed freely within these populations, but not across these populations. 2 Individuals, Assemble! Theories ofSpeciation.... Allopatric Speciation Allopatric speciation (from the ancient Greek allos, "other" + Greek patra, "fatherland") occurs when a population is split into two geographically isolated populations, such that they are able to interbreed freely within these populations, but not across these populations. Populations often exist across separate territories or distributed across the extent of their habitat. But, they can consistently interbreed by either traveling between territories or by meeting up with members of other territories during yearly migration. By continuing to interbreed, the gene pool maintains a spread consistency, and evolution occurs only as traits are naturally selected across the entirety of the population. 2 Populations often exist across separate territories or distributed across the extent of their habitat. But, they can consistently interbreed by either traveling between territories or by meeting up with members of other territories during yearly migration. By continuing to interbreed, the gene pool maintains a spread consistency, and evolution occurs only as traits are naturally selected across the entirety of the population. But the face of the earth is always changing. Depending on the species and its territory, a toplogical shift such as a river swinging its course, the rise of a mountain range, climate shift such that a species depended on for food or other resources splits its own territory, or any other spontaneous reallignment of biome and geography could potentially rend a population into two halves, which are unable to continue interbreeding as a result. Separated in this way, the two populations will take their separate courses in evolution. It is now highly unlikely that, without interbreeding to normalize the distribution of genetic traits across the entire population, these two populations will continue as a single species. The vicissitudes of their territories and the separate ecosystems means that no matter how similar, some mutations' advantages will increase fitness in one area, and not in the other. Over time, natural selection will effect changes in the population's genetics such that they become separate species, despite their common genetic heritage. Cases ofAllopatric Speciation - The variation between populations on different islands in a chain, such as on the Galapagos, shows how evolution proceeds in its own direction within a particular habitat. The species called "Darwin's finches" is a good example ofthis, though Darwin himselfonly connected them to the process ofevolution later, after he returned to England. - The African elephant, once thought ofas a single species, is now considered as three subspecies by some researchers. Geographic isolation is theorized as creating these differences. The Asian elephant also has four distinct sub-species. - Fruit flies can show mating preference based on food source, after being isolated to a particular food source for only eight generations. But the face of the earth is always changing. Depending on the species and its territory, a toplogical shift such as a river swinging its course, the rise of a mountain range, climate shift such that a species depended on for food or other resources splits its own territory, or any other spontaneous reallignment of biome and geography could potentially rend a population into two halves, which are unable to continue interbreeding as a result. Separated in this way, the two populations will take their separate courses in evolution. It is now highly unlikely that, without interbreeding to normalize the distribution of genetic traits across the entire population, these two populations will continue as a single species. The vicissitudes of their territories and the separate ecosystems means that no matter how similar, some mutations' advantages will increase fitness in one area, and not in the other. Over time, natural selection will effect changes in the population's genetics such that they become separate species, despite their common genetic heritage. After speciation has occurred, the species will live independently even if their territory is rejoined. Sexual selection or genetic and physiological incompatibilities will ensure that the gene pools of the two species are separate from now on. The two populations are now species sharing territory, or if evolved far enough apart, they might occupy different niches as if they were separate territories. After speciation has occurred, the species will live independently even if their territory is rejoined. Peripatric Speciation 3 Theories ofSpeciation.... Peripatric speciation is a subform of allopartic speciation. The main principle is the same, but it occurs with a much smaller sub-population. Peripatric speciation begins when a small portion of the species separates from the main population. This separation typically occurs over a distance, as this sort of geographic separation is more likely to happen to a small number, whereas a territorial split would take a sizeable amount of the population with it. d d+1 d+2 d+3 d+x Peripatric Speciation Peripatric speciation is a subform of allopartic speciation. The main principle is the same, but it occurs with a much smaller sub-population. Once the population has a chance to establish itself, it can become its own population, growing to fill this new territory. Because of the small initial size of the population, this is a characteristic situation for genetic drift. "Founder's Effect" is a loss of genetic variation occurring when a small number of individuals increases into a large population. This loss of variation can cause the propagation of rarer traits. Once the population has a chance to establish itself, it can become its own population, growing to fill this new territory. Examples ofFounder's Effect - Island Populations, such as the human populations ofIceland, the Faroe Islands, Easter Islands, and Pitcairn Islands all have relatively low genetic variation. - There is a statistically high population ofdeafpeople born on Martha's Vinyard. Additionally, this resulted in the development ofMartha's Vinyard Sign Language. - Polydactyly (extra fingers and toes) are more common in endogamous Amish populations than the general population. Because of the small initial size of the population, this is a characteristic situation for genetic drift. "Founder's Effect" is a loss of genetic variation occurring when a small number of individuals increases into a large population. This loss of variation can cause the propagation of rarer traits. - After a typhoon killed all but twenty inhabitants on the island ofPingelap, total color-blindness in the since-regained population is at 5%. Examples ofFounder's Effect - Island Populations, such as the human populations ofIceland, the Faroe Islands, Easter Islands, and Pitcairn Islands all have relatively low genetic variation. - There is a statistically high population ofdeafpeople born on Martha's Vinyard. Additionally, this resulted in the development ofMartha's Vinyard Sign Language. - Polydactyly (extra fingers and toes) are more common in endogamous Amish populations than the general population. In such a bottleneck as when a population's genetic diversity is shifted by Founder's Effect, peripatric speciation can occur. It is essentially the same as the Founders' atypically large genetic drift affect on the gene pool of the population, but with adaptive mutation to a point at which the smaller population cannot interbreed with the main population any longer. - After a typhoon killed all but twenty inhabitants on the island ofPingelap, total color-blindness in the since-regained population is at 5%. At this point, if members of the departing population were to return to the territory of the original population, they would find themselves to be two separate species, inhabiting the same territory. Or, two overlapping territories. The dynamics of competition and cooperative survival of the gene pool are the territory, in that they define niches within the system of resources, and thusly, the entirety of the individuals' lives of breeding and survival. In such a bottleneck as when a population's genetic diversity is shifted by Founder's Effect, peripatric speciation can occur. It is essentially the same as the Founders' atypically large genetic drift affect on the gene pool of the population, but with adaptive mutation to a point at which the smaller population cannot interbreed with the main population any longer. d d+1 d+2 d+3 d+x Examples ofPeripatric Speciation - The Australian Pacific Robin - Fruit flies have been induced to form separate reproductive populations as as result ofbottlenecking - Polar Bears are believed to have diverged from the Brown Bear species, after straying out from the margin ofthe parent species' range. At this point, if members of the departing population were to return to the territory of the original population, they would find themselves to be two separate species, inhabiting the same territory. Or, two overlapping territories. The dynamics of competition and cooperative survival of the gene pool are the territory, in that they define niches within the system of resources, and thusly, the entirety of the individuals' lives of breeding and survival. Theories ofSpeciation.... Parapatric Speciation 4 Follow the territory, species! Find your niche! Theories ofSpeciation.... Parapatric Speciation 4 Parapatric speciation, or the formation of a ring species, is a unique and more complicated pattern. Variations in mating habits among members of a species in a continuous, but gradiated habitat prevent gene flow across the entire species, and create a continuum of speciation in small steps, wherein neighbors can interbreed or hybridize with neighbors, but the entirety cannot. This sort of speciation happens in unique terrain, where the territory of the species stretches over a thin span of space, and slight variations make the species' niche different along its length, and yet still conducive to that species' survival. Parapatric speciation, or the formation of a ring species, is a unique and more complicated pattern. Variations in mating habits among members of a species in a continuous, but gradiated habitat prevent gene flow across the entire species, and create a continuum of speciation in small steps, wherein neighbors can interbreed or hybridize with neighbors, but the entirety cannot. This sort of speciation happens in unique terrain, where the territory of the species stretches over a thin span of space, and slight variations make the species' niche different along its length, and yet still conducive to that species' survival. There is no geographic boundary line in parapatric speciation, but a geographic gradient. Because of convenience, as well as shared specialization to deal with their local conditions, members of a population are more likely to breed with those near each other, and not those further away. Accordingly, genetic variations will build up in zones along this gradient, reinforced by these breeding choices. There is no geographic boundary line in parapatric speciation, but a geographic gradient. Because of convenience, as well as shared specialization to deal with their local conditions, members of a population are more likely to breed with those near each other, and not those further away. Accordingly, genetic variations will build up in zones along this gradient, reinforced by these breeding choices. When this gradient rejoins itself, a ring species results. This can happen around a ring of islands, around a pole, or even around elevations of valleys. When this gradient rejoins itself, a ring species results. This can happen around a ring of islands, around a pole, or even around elevations of valleys. Larus gulls are a prime example of a ring species. Seven different subspecies extend in a ring around the Artic circle, each able to interbreed or hybridize with its neighbors in a circular continuum except for one pair, the "gap" in the ring, so to speak. A separate species has resulted from the overlapping ends of a gradient genepool. Other ring species include the Ensatina salamanders, which encircle the Central Valley in California, and the Greenish Warbler, which ring the Himalayas. Larus gulls are a prime example of a ring species. Seven different subspecies extend in a ring around the Artic circle, each able to interbreed or hybridize with its neighbors in a circular continuum except for one pair, the "gap" in the ring, so to speak. A separate species has resulted from the overlapping ends of a gradient genepool. Theories ofSpeciation.... 5 Sympatric Speciation Allopatric and peripatric speciation have rigid geographic boundaries that encourage population separations that result in speciation. Parapatric speciation does not have a barrier so much as a gradient, that spreads the obstacles to complete interbreeding between members of a population over a varied terrain, thereby creating a varied speciation. With sympatric speciation there is a firm population division. However, there is no geographic boundary. This type of speciation occurs within a single population in a single territory, but is no less of a genetic split resulting from a bifurcating gene pool. One might wonder, if this sort of speciation can occur, on what basis can we say that a population is a single population, or that it inhabits a single territory? What sorts of topological segmentation is there that is not geographic, and yet no less affects the breeding behavior of a species? Sympatric Speciation Allopatric and peripatric speciation have rigid geographic boundaries that encourage population separations that result in speciation. Parapatric speciation does not have a barrier so much as a gradient, that spreads the obstacles to complete interbreeding between members of a population over a varied terrain, thereby creating a varied speciation. With sympatric speciation there is a firm population division. However, there is no geographic boundary. This type of speciation occurs within a single population in a single territory, but is no less of a genetic split resulting from a bifurcating gene pool. One might wonder, if this sort of speciation can occur, on what basis can we say that a population is a single population, or that it inhabits a single territory? What sorts of topological segmentation is there that is not geographic, and yet no less affects the breeding behavior of a species? Within a population group, slight variations begin to occur, which exhibit traits with behavioral or physiological differences that are perceivable, at least to the extent that it affects breeding behavior. For example, certain members of a population might be taller or shorter, or have markings that are slightly different. Within a population group, slight variations begin to occur, which exhibit traits with behavioral or physiological differences that are perceivable, at least to the extent that it affects breeding behavior. For example, certain members of a population might be taller or shorter, or have markings that are slightly different. If the trait encourages sexual selection in such a manner that it isolates particular individuals from the wider population, it becomes its own limiting effect on the continuity of the overall gene pool. If, to continue the example, tall individuals only wish to breed with tall individuals, and this choice is widespread enough, eventually, the genotype expressing that trait will become withdrawn from the general population and form its own subpopulation. The border in this case does not physically exist, but is imposed topologically on the breeding environment by the population's behavior. And yet, these behaviors, when determining sexual selection, are just as real to the gene pool as the geographic obstacle of a mountain or a river. If the trait encourages sexual selection in such a manner that it isolates particular individuals from the wider population, it becomes its own limiting effect on the continuity of the overall gene pool. If, to continue the example, tall individuals only wish to breed with tall individuals, and this choice is widespread enough, eventually, the genotype expressing that trait will become withdrawn from the general population and form its own subpopulation. The border in this case does not physically exist, but is imposed topologically on the breeding environment by the population's behavior. And yet, these behaviors, when determining sexual selection, are just as real to the gene pool as the geographic obstacle of a mountain or a river. Sooner or later, a separate species results, living among the original species. What was begun as a matter of sexual choice is reinforced by genetics, that has had a chance to mutate within the subpopulation, and via natural selection and genetic drift permanently branch from the main population. Examples ofSpecies That Have Branched Due to Sympatric Speciation - Cichlids ofLake Nabugabo in East Africa - Three-spined sticklebacks in British Columbia - The Hawthorn fly in North America Sooner or later, a separate species results, living among the original species. What was begun as a matter of sexual choice is reinforced by genetics, that has had a chance to mutate within the subpopulation, and via natural selection and genetic drift permanently branch from the main population. Examples ofSpecies That Have Branched Due to Sympatric Speciation - Cichlids ofLake Nabugabo in East Africa - Three-spined sticklebacks in British Columbia - The Hawthorn fly in North America Speciation: A Diagram Book designed and printed by Adam Rothstein for Bordertown Information in the text is sourced through Wikipedia. Please visit the article on "Speciation" as well as relevant sub-articles for the latest word on speciation theories, as well as the scientific research status of particular species as regards these theories. Speciation: A Diagram Book by Adam Rothstein is licensed under a Creative Commons AttributionNonCommercial-ShareAlike 3.0 Unported License. http://www.poszu.com/projects/speciation 201 1
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