Speciation Reading: Freeman, Chapters 25-26 Many of the figures, and most of my examples, are from Douglas Futuma’s Evolutionary Biology, an excellent reference if you want to know more about speciation Species Concepts • Biological – “Species are groups of actually or potentially interbreeding individuals that can mate and produce fertile offspring”-idea was promoted by Ernst Mayr, an evolutionist who worked on birds. • In practice, this applies to most species, but in many cases, it is simply impossible to test whether two species have the potential to interbreed. • Phylogenetic – A species is a lineage, separate from other such lineages, perpetuated ancestor to descendant, over time. • Morphological – Morphological criteria are used to define species. Clearly, each school of thought has its strengths and its drawbacks. The biological species concept is great conceptually, but virtually impossible to put into process in most situations. Fossil species, species which do not reproduce in the lab or zoo (the vast majority), and asexual species are sticky issues. The morphological species concept is what most taxonomists actually use in practice, because it is expedient, but it has many disadvantages. It is subjective. Cryptic species, which look identical to humans but are in fact reproductively isolated, are problematic. The phylogenetic species concept is becoming increasingly used-it is most useful when the scientist has a very clear idea what type of “lineages” they are looking at. Speciation • The theory of evolution must explain the origin of new organisms. From the beginning, the origin of species has been a focal point of evolutionary theory. – Ironically, Darwin was wrong about the origin of species. He assumed that, given enough time, natural selection would inevitably produce them. This is not actually the case. It takes reproductive isolation. – Macroevolution is the origin of new taxonomic groups. – Speciation is the origin of new species. With extinction, it is one of two keystone processes of macroevolution. Cladogenesis • Cladogenesis, the origin of lineages, is the budding of a new species from a parent species that continues to exist. • Cladogenesis promotes biological diversity by increasing the number of species. • Although it culminates over thousands or millions of years, cladogenesis is a real event. New species originate by cladogeneis, which is ultimately responsible for the origin of every major group of animals. Why Do We Have Separate Species At All? • Sympatric species live in the same place. Without some mechanism preventing allele and gene exchange among sympatric species, distinct species would be impossible, we would probably see a continuum from one form of life to another. • Barriers to allele flow are called isolation mechanisms. • Isolation mechanisms allow sympatric species to exist. • Without isolation mechanisms, closely related species would hybridize: allele flow and recombination would eventually transform them into a single, polymorphic species. The Evolution of New Species Results From (and also causes) Barriers to Allele Flow. • At the time when a genetic or behavioral mechanism evolves that keeps populations of a species from interbreeding, two new species are formed. • As we shall see, this is frequently the result of a geographic barrier, although it may be the result of a chromosomal change or habitat preference. There are Two Basic Categories: Prezygotic Isolation Mechanisms Postzygotic Isolation Mechanisms • Presygotic isolation mechanisms prevent mating, so that gametes of sympatric species never form hybrid zygotes. • Postzygotic isolating mechanisms act after a mating has occurred, to prevent fertilization or to prevent potential hybrids from passing on their genes. Prezygotic Isolation Mechanisms • • • • • Habitat Isolation Temporal Isolation Behavioral Isolation Mechanical Isolation Gametes Die Habitat Isolation • Habitat isolation occurs because sympatric species meet due to differences in their habitat preference. • Examples of habitat isolation: • Sympatric species of spadefoot toads (Scaphiopus) seldom meet because they prefer different soil types. • Many species of closely-related parasites, such as bird lice, never meet because they live and mate on different hosts. Temporal Isolation • Temporal Isolation: Occurs because species mate at different times. • Examples: • Different species of plants frequently have differing flowering seasons. • Closely related species of fireflies frequently mate at different times of night. Behavioral Isolation • Behavioral, or Ethological isolation mechanisms include differences in courtship behavior, differences in chemical signals or vocalizations, and differences in color or morphology that allow individuals to recognize their own species. • They are a very common mechanism keeping closely-related sympatric animals from interbreeding Examples: • Female fireflies respond only to the light pattern emitted by their own species. Sympatric species of fireflies emit different light patterns. • Experiment: Gulls normally mate only with their own species: artificial hybridization can be induced by modifying the contrast in color between the eye and the face. Thus, the isolation mechanism is behavioral. Mechanical Isolation • Mechanical isolation occurs because the sexual organs of closely-related sympatric species are incompatible: they do not fit together. • This is thought to be an important isolation mechanism in arthropods, particularly insects and millipedes. Sperm of various mammals Gametic Mortality Gametes are frequently very specialized cells, which can only perform well in the reproductive tract of the opposite sex of the same species. In many angiosperms, for instance, pollen transferred to the stigma of another species will not germinate, or if they do, will not form a pollen tube. Postzygotic Isolation Mechanisms • • • • Zygote dies after fertilization Hybrid Inviability Hybrid Sterility Low Hybrid Fitness Hybrid Inviability • Some species that do not ordinarily interbreed occasionally do so. Frequently, the progeny of these interspecific matings die at some point during their development. • Example: Hybrids between the frogs Rana pipiens and Rana sylvatica do not survive more than a day or so. Hybrid Sterility • Hybrid Sterility occurs when the hybrid of an interspecific mating is unable to reproduce. • Examples: Mules are the hybrid of a horse and a donkey, they do not form normal sperm. • Hybrids between Drosophila melanogaster and D. pseudoobscura have atrophied testes and are sterile, Low Hybrid Fitness • If interspecific hybrids do survive, they often have very low fitness, this effectively keeps them from spreading genes from one of their parent species to the other parent species. • Example: Dog-Wolf hybrids are perfectly viable, but they are considered to be unsuitable pets in most areas. • Wild wolf populations do not accept hybrids, they are killed on sight. There Are Two Basic Types of Speciation: Allopatric and Sympatric • Allopatric Speciation: Involves a geographic barrier. • Sympatric Speciation: Does not involve a geographic barrier. Allopatric Speciation • Allopatric speciation involves a geographic barrier that physically isolates populations of a species and blocks gene flow. • Once isolated, allopatric populations (living in different places) accumulate genetic differences due to natural selection, genetic drift, and new mutations. If the Geographic Barrier is Removed, the Two Species May: • 1) meld together by allele flow and recombination to once again form a single species. • 2) remain reproductively isolated. • The fate of the new, incipient species, depends upon whether isolation mechanisms have evolved during the period of isolation. • These isolation mechanisms may be premating or postmating. • Premating isolation mechanisms may evolve in incipient species that have postmating isolation, to reduce the probability of incorrect matings and the subsequent loss of fitness. Example of geographic isolation and possible allopatric speciation: Two closely-related species of antelope squirrels live on opposite sides of the grand canyon. On the South rim is Ammospermophilus harrisi, on the North rim is Ammospermophlus leucurus. Birds, and other species that can cross the canyon, have not diverged into different species on opposite sides. Example: the Drongo • The Drongo is a black bird with a crest of feathers, it is highly variable in behavior and appearance throughout its range. • Each semi-isolated population has its own appearance. • A “double” invasion probably occurred in Tasmania over the course of the past few thousand years. • The species Acanthiza pusilla is widespread on the Australian continent. Tasmania has a slightly differentiated population of this bird. • Another, reproductively isolated species, A. ewingi, that is even more differentiated from Australian Drongos in appearance and morphology also inhabits Tasmania. • During the last ice age, when sea level was lower, Tasmania was part of island, this is probably when the ancestors of A. ewingi invaded the island. Eventually they evolved reproductive isolation from their Australian counterparts. • When A. pusilla re-invaded the island more recently, the two species were able to co-exist because they are reproductively isolated. Adaptive Radiation on Islands ·Island chains frequently produce many new species. ·Islands chains provide barriers that facilitate invasion and re-invasion by different species. ·This is the probable mechanism for the proliferation of Darwin’s finches on the Galapagos. ·The Hawaiian islands once supported thousands of unique Drosophila flies that probably evolved by a similar mechanism How Long Does Allopatric Speciation Take? • Nobody knows for sure, and it depends upon the group. – McCune and Lovejoy, based on a study of reproductive isolation in 40 pairs of allopatric fishes, estimated that, for fishes, it takes between .8 and 2.4 million years for reproductive isolation to evolve. • Hurt and Hedrick conducted interesting studies of incipient allopatric speciation in the Sonoran topminnows – Poeciliopsis sonorensis and Poeciliopsis occidentalis occupy different river drainages in Arizona http://www.nativefish.org • Based on molecular evidence, the two species/subspecies have been isolated for between one and two million years. • McCune and Lovejoy found that males of each species prefer to mate with females of their own species, but given a choice, they will hybridize. • There was evidence of reduced hybrid fitness, especially when hybrids were crossed with one of the original species. – Brood sizes were smaller, and there was an unusual, male-biased sex ratio for these crosses. • Are they separate species? – Clearly, the answer is subjective. In this case, if the populations were to mix together, the process of reinforcement would probably complete the job….so probably yes. – Reinforcement-natural selection on females of these incipient species pairs to avoid mating with males of the wrong species, thus avoiding the cost of producing unfit, hybrid offspring. Sympatric Speciation • Sympatric speciation results from intrinsic factors, such as chromosomal changes and nonrandom mating. • Sympatric populations become genetically isolated even though their ranges overlap. Mechanisms of Sympatric Speciation • Polyploidy: allopolyploidy and autopolyploidy • Nonrandom mating: I.e., host shift Polyploidy and Interspecific Hybridization • Polyploidy: Disorders of meiosis cause the accidental formation of gametes that are 2N rather than N. Union of two of these gametes produces a zygote that is 4N. The chromosome number has doubled, instantaneously producing a potential new species! This produces an autopolyploid. • Interspecific Hybridization: Gametes of two species meet and form a hybrid set, usually the hybrid set is sterile, but sometimes it is not. This produces an allopolyploid, which is usually infertile (the chromosomes can not pair during meiosis), but may reproduce asexually. If the chromosomes double by a disorder of meiosis, it produces a potentially sexual species • These mechanisms are probably common in plants, but rare or absent in animals. • Plants are frequently capable of self-fertilization, and some can survive with double the normal number of chromosomes, and most can propogate asexually. • Many common agricultural plants are the products of one, or both of the mechanisms above. • Examples: Wheat is doubly autopolyploid, the product of an interspecific hybridization, then a doubling of chromosomes, then another interspecific hybridization, then another doubling of chromosomes, to produce a fertile hexaploid. • Yellow bananas are allopolyploid, the product of two interspecific hybridization events. Host Shift • Nonrandom mating due to host shift: Many species of parasites mate on or nearby the host. A shift to a new species of host therefore reproductively isolates the parasites exploiting the different species of hosts. • Example, the apple maggot: The best studied example of this occurs in the apple maggot. Apple trees are not native to the US, they were introduced here in the nineteenth century. Following their introduction, the hawthorne maggot began to feed on apples. The flies cue in to the smell of their original host, so apple and hawthorne maggots are now reproductively isolated: and considered to be different species. Parapatry • Speciation is not always clear-cut, there are many examples of SOME populations being reproductively isolated, while OTHERS are able to interbreed. This is sometimes called Parapatry. • Example: A well known example involves California garter snakes, Thamnophilus sp.. • Each population of land snakes is able to interbreed with the populations closest to it, but not with more distant populations. In some cases, however, a distant population has come back around to encounter a distantly related population. In these cases, they do not interbreed. A similar relationship exists for water populations. Some water snakes can even interbreed with the local land snakes These figs are from Futuma’s Evolutionary Biology Phylogeny, Taxonomy, and Systematics • Phylogeny: The phylogeny of a group is a “family tree” describing how species are related. The branching pattern of different groups of organisms is caused by repeated cladogenesis. • Systematics is the study of phylogeny. • Taxonomy: Is the process of describing and naming organisms. Our modern process of taxonomy is based on phylogeny, so an understanding of phylogenetic relationships-names reflect relatedness. The Taxonomic Heiriarchy • • • • • • • • • • Every species has a place in the taxonomic heirarchy: Human Mud-Dauber Species Homo sapiens Trypoxylon politum Genus Homo Trypoxylon Family Hominidae Sphecidae Order Primates Hymenoptera Class Mammalia Insecta Phylum Chordata Arthropoda Kingdom Anamalia Anamalia Domain Eukarya Eukarya Inferring Phylogeny • The branching pattern of a phylogenetic tree reflects its ideal place in our taxonomic hierarchy. • Classification schemes are hypotheses of past history based on the available evidence. • Like all hypotheses, they make predictions that can be tested by future study. The phylogeny of a group of organisms can be inferred from the following lines of evidence • Shared characteristics passed down from an ancestor, called homologies. – These are essential in inferring a phylogeny. • Morphology and DNA sequences are very useful places to look for homologies • Biogeography • The Fossil Record Homology • A character state is homologous in two species when it is inherited by both from a common ancestor. • The most widely accepted school of systematics today is called cladistics. • Cladistics infers the pattern of phylogeny based on homologies. Groups are constructed based on shared characteristics inherited from a common ancestor, that no other group has. • Trees are constructed by creating a nested series of such groups. Types of Characters Used in Cladistics • Apomorphy-an evolutionary novelty for a group. • Plesiomorphy-an evolutionarily primitive state • Synapomorphy-a novel (derived) trait that a group has inherited because the common ancestor of that group had a novel characteristic and passed it on. • Synplesiomorphy-an evolutionarily primitive trait that a group has inherited because the common ancestor of that group had inherited the primitive condition, unchanged, from an earlier group. • Note that these terms are relative-a synplesiomorphy for one group may be a synapomorphy from the larger group it came from. Example of a Homology which is a Synapomorphy Homoplasy • If a character has evolved more than once, if possessed by two species but not present in the common ancestor, it is called a homoplasy. • One form of homoplasy is called convergent evolution, it is quite common because different species are often subject to similar selective pressures. • Homoplasy, when mistaken for homology, can obscure the pattern of evolutionary history. Example of a Homoplasy-not a homology Extinction 99.9999% of all species that have ever lived are extinct • Extinction has always been a major force in macroevolution. • Species do not last forever…the mean expected lifespan of a marine bivalve is about 14 million years, and the mean expected lifespan of a terrestrial mammal might is about a tenth that. • Climate change, natural disasters, and other phenomena have always caused extinction. • As some species originate, they inevitably drive other species extinct. • Extinctions, in turn, pave the way for speciation. • An adaptive radiation is a wave of speciation that occurs as a new habitat is colonized by a lineage, or in the wake of the extinciton of another lineage. • An adaptive radiation of mammals followed the extinction of the dinosaurs. Background Extinction • At all times in history, groups of organisms have a “background extinction rate”…species go extinct because of normal ecological or evolutionary processes, and as they disappear, other species take their places. – For instance, on oceanic islands, the arrival of a predator, such as a monitor lizard or a snake, might precipitate the extinction of ground-nesting birds. Any such birds that are endemic to the island (that is, they live nowhere else) are gone for good. – But oceanic islands come and go, as geological forces shape the Earth’s crust, and such extinctions are considered to be “normal”. Mass Extinctions • The history of life on Earth has been punctuated by a series of mass-extinctions. – Extinction rates are much higher than background rates for a short period of time. – Some are better understood than others, but they have profoundly influenced the evolution of life on Earth. – Over the last 500 million years, there have been several major mass extinctions (not counting the current one): – here are some big ones • • • • • 1) The Late Devonian 2) Mid-Ordovician 3) Permian-Triassic 4) Late Triassic 5) Cretaceous-Tertiary. The Human Mass-Extinction • Since the development of agriculture, 10,000 years ago, humans have modified an increasing proportion of the Earth’s resources for our own purposes. • Humans impact has caused extinction rates to be 10 to 1000 times greater than any time in the last 100,000 years. • For example-one estimate for the recent background extinction rate for birds is one species extinction per 400 years. – If only this natural rate of loss affected the number of bird species, no more than a couple of extinctions should have occurred in the past 800 years. – Scientists estimate that the actual loss during this time period lies somewhere between 200 and 2,000. • We have set in motion a mass extinction, one of the largest, that will not culminate until thousands of years from now. • Humans have extensively modified the biosphere • The human population passed 6 billion in the year 2000, and is growing at a rate of almost 2% per year. • Each human uses so much energy and so many resources that our activities influence virtually every aspect of the biosphere. • In temperate areas, nearly all the land area that is suitable for agriculture is plowed or fenced. • Worldwide, more than 35% of all land area is used for farms or permanent pastures. Much of the rest is grazed or logged on a regular basis. From 35-45% of Global Net Primary Productivity now goes to serve human needs. • In aquatic ecosystems as well, an increasing amount of productivity is harvested by humans. Nearly every major fishery in the Northern Hemisphere has showed strain from overharvesting, and many have collapsed. Major sources of anthropogenic extinction • Habitat destruction and habitat fragmentation • Habitat Change and Disruption of Ecosystem Processes • Introduction of Exotics • Overexploitation Habitat destruction and habitat fragmentation • Most of the grasslands and forests of the Northern Hemisphere were destroyed by the end of the nineteenth century, the grasslands of the southern hemisphere are now vanishing, and tropical forests are disappearing at a rate of about 2% per year. This type of destruction has become the norm for most biological communities, as the human population expands our economic needs require resources from more and more land. The remaining habitat is often broken into many small fragments, which are separated by large areas of land under cultivation or other human uses, effectively reducing a single "continent" into many "islands". Fragmented Habitats Support Smaller Populations • Essentially, every habitat fragment becomes a biological "island" (analogous to continental shelf islands, rather than the oceanic kind). – As in the Mac Arthur Wilson model, the smaller the island, the smaller the population of any given species it can support. – Small populations are at much greater risk of extinction due to random events, such as weather, disasters, and natural fluctuations in their population and sex ratio. This part of Canada used to be a continuous swath of natural communities. Here is a fragment seen from the air • Additionally, smaller populations support less genetic variation, which could lead to the fixation of harmful alleles and the ultimate extinction of the population (for very small fragments), or simply inhibit their ability to evolve in response to changing conditions Fragmented Habitats Frequently Lack Critical Ecosystem Processes • Edge effects fundamentally alter habitat. For certain species, this can be critical to their ability to survive. For instance, places where human habitation borders nature preserves frequently have weedy plants, fire is controlled, domestic cats and dogs escape and prey on native wildlife, and human noise and activity disturb the behavior of certain animals. • The edge habitats have different effects on different species. – Some large mammals, such as coyotes and raccoons, reach much higher densities in edge habitats because they are able to take advantage of human resources (garbage), and return to the safety of the preserve. • Taking this a step farther,raccoons in the US, and red foxes in England, have even penetrated urban areas to become part of the city, reaching high densities. – Other mammal species cannot tolerate edge environments, wolves and mountain lions do not like humans and cannot live on the edge (in cases where they try, very bad things might happen. This is a natural area in Massachusetts. • Example: The brown headed cowbird is a native to the United States. It is a brood parasite, evicting the eggs of other species to replace them with its own. Cowbirds prefer edge habitats. Now that forests are fragmented, there are few safe areas from cowbirds, and forest interior species such as bluebirds are suffering a major loss of fitness in some areas. Habitat Change and Disruption of Ecosystem Processes –Surviving areas of natural habitat often change because humans have fundamentally altered natural ecosystem processes. Examples – Ladys' Slipper Orchids. There are probably about 25, 000 species of orchids worldwide, and they are being lost faster than they can be classified. – Orchids are typically tightly coevolved in mutualistic relationships with other species, and the loss of any of these relationships can lead to extinction. • Ladys' slippers are a very diverse group that occupy a wide variety of habitats in the Northern Hemisphere. They are in decline even in protected areas, such as Indiana Dunes. • Human activities have altered their ecosystems. Ladys' slippers need a mutualistic fungus to germinate and grow for the first few years. Airborne nitrogen compounds (mostly from automobiles) effectively "fertilize" vast areas of ground and may put the mutualistic fungi at an ecological disadvantage. • Also, the widespread application of pesticides, the human tendency to groom and "clean up" areas of open sand and fallen wood, and the introduction of the honeybee to North America have caused the Andreneaid bees that would normally pollinate these plants to disappear from many areas. – Pacific Salmon are very important ecologically and economically off the West Coast of North America. – Salmon species have experienced dramatic declines over the past few decades due to a variety of factors, many of which result from human habitat modification. – Hydroelectric dams have resulted in increased juvenile mortality and made many habitats inaccessible to migrating salmon. – Additionally, human logging and agriculture has silted and modified many of their upstream habitats, causing a drop in recruitment. Introduction of Exotics • Human activities are creating the worldwide equivalent of the "Great American Faunal Interchange". This is an uncontrolled experiment in community ecology, with the potential result of a massive loss of gamma diversity worldwide caused by the loss of endemic species. Zebra Mussels • In 1998, the zebra mussel was discovered in Lake St. Claire near Detroit. It was introduced to the Great Lakes from the Caspian Sea, probably in the ballast water from a cargo ship, sometime around 1985. This mode of dispersal is very common, in 1982 the comb jelly (a ctenophore) was introduced to the Black Sea in a similar manner. Comb jellies increased in number until they amounted to an estimated 90% of animal biomass in the Caspian. • They have since spread throughout the Great Lakes • • • • region and throughout the Mississippi and Ohio River valleys. It forms dense clusters of individuals, and can clog the water intakes of electrical power stations, water stations, and other industrial facilities. Zebra mussels are incredibly effective filter feeders. Zebra mussels actually make the water much clearer, but alter native communities of organisms in the process. In the Hudson River, phytoplankton biomass decreased 85% after zebra mussels invaded, zooplankton decreased 70% as a result. The zebra mussel is a very effective competitor. Extinction of native bivalves will almost certainly result from this introduction. You may have noticed that Chicago water tastes weird during the summer, that is because the water is now clean enough to allow the growth of cyanobacteria deep enough in the lake to be pulled into the water intakes. The residue of cyanobacteria toxins has an off taste. Honey Bees • Honey bees are native to Europe and Asia. Apis mellifera, is a European species that is widely cultivated for honey, beeswax, and as a pollinator. European immigrants probably introduced the honeybee to North America in the nineteenth century (Native Americans called it "white man's fly".) It is a very effective competitor, and displaces native bee species. • Recently, honeybees themselves have taken a hit, when the varolla mite was introduced in the 1980's. The overuse of insecticides, and widespread destruction of habitat, have decimated North American bee populations, both native and non-native. – The Snake that Ate Guam. Boiga irregularis, the brown tree snake, is an arboreal snake native to New Guinea, Australia, and the Solomon Islands. It is a small, nocturnal, rear-fanged snake. – Boiga irregularis was introduced to Guam in the late 1940s, probably by hitching a ride in the wheel well of a plane. Since that time, it has literally eaten most of the endemic birds of Guam to extinction. Since there are no other native snakes in Guam (other than a blind, burrowing species), the bird fauna there evolved no natural defenses. • Thus, is an incredibly effective predator of birds and their nests. – In Australia, competition and predation keep it in check, but the simpler ecosystem of Guam has allowed it to increase in numbers to up to 20 individuals per square acre of jungle (among the highest ever recorded for a snake). – It also causes other problems in Guam, including numerous power outages resulting from large numbers of snakes resting on power lines. Garlic Mustard, Purple Loostrife, Multiflora Rose – These are three more cases of an introduced species being too good at what they do. All three plants were introduced intentionally in the nineteenth century. Each of the three has become so common that it is likely to displace other species. For example, in some East Coast marshes, purple loosestrife amounts to 90% of the vegetation, displacing native sedges and other plants. Overexpolitation • Stellar's Sea Cow-this huge sirenian mammal lived in the reached a length of 26 feet and could way seven thousand pounds or more. It existed on a diet of kelp, and could not dive or swim quickly. • It was delicious, and was hunted to extinction by sailors within 30 years of its discovery What Makes A Species Vulnerable to extinction? • • • • • Endemism Rarity Small Population Size Ecological Specialization Beauty/Usefulness to humans/Competitor with Humans • Endemism- Species that are restricted to a particular, small area, are more vulnerable to extinction • Rarity-Rarity is not the same thing as endeminsm, endemics can be very common in the restricted area where they do occur. "Naturally rare" species have low population densities, but may be widely distributed and have respectable population sizes. We do not completely understand the ecological factors that make some species "naturally rare", but when a common species gradually becomes rare, it is often a prelude to extinction. "Naturally rare" species can be a challenge to conservation, because they are difficult to monitor and it is very difficult to ensure that sufficient habitat is set aside for them. • Small Population Size-Small population sizes render a species very vulnerable to extinction, through reduced genetic variation via genetic drift, the potential for inbreeding depression, demographic stochasticity caused by random ecological disasters and, for sexual species, the small chance that every individual in the population might be born the same sex. • Ecological Specialization-Ecological specialists are more prone to extinction because there are only a few ways they can 'fit themselves into" an ecosystem. They must have certain interspecific relationships in order to feed, obtain mates, have places to live, or maintain competitive superiority. The loss of other species in the community, or habitat change due to human activity, can change these factors, and render a formerly successful species vulnerable to extinction. • Useful to Humans or A Competitor of Humans-Humans have a way of killing all the pretty things, harvesting all the useful things, and hunting to extinction everything that could be perceived as a competitor. For instance, fishermen in San Francisco are prone to despising the California Sea Otter, despite its important place in the ecosystem of the California Coast, because of its status as a competitor. They are protected now, however, they were nearly hunted to extinction for their pelts. Species that cross the paths of humans sometimes suffer for it. • Economic Considerations • Any conservation plan that does not take human economics into account is prone to failure. It is very difficult to set aside a habitat and protect it from all human activity. The closest we have ever come are on military bases and nuclear test sites (the conservation effect was unintentional at first), and some private organizations (Nature Conservancy) buy natural land and simply fence it off. Even these exceptional preserves have neighbors, and are occasionally eyed by developers and government reclassification. • The vast majority of preserves must balance the needs of human ecotourists, indigenous peoples, neighbors, and government budget considerations against conservation goals. The Future of Our Own Species – It is one of the strange ironies of our existence that, though the actions of our species modify the biosphere to an extent unprecedented in the history of the earth, as individuals, we do not necessarily feel any collective responsibility for our actions. – The future of our own species will depend, to a very large extent, upon decisions we make as individuals, regarding our priorities. It is quite possible for our species to survive for many thousands of years more, but this is likely only if this generation takes additional steps to ensure that the planet will remain habitable to our own species.