Kalafsky 1 Mrs. Kalafsky - AP Biology Ecology Summer Assignment 2013 Welcome to AP Biology! In an effort to be prepared for the AP Biology exam, we have a tremendous amount of work ahead of us before May 12, 2014! I hope you are all up for the challenge! To start us off I have attached your 2014 Summer Assignment. Books: It is imperative that you order your text with Mastering Biology immediately. Use of the text is required for the summer assignment. Lack of a text will not be accepted as an excuse for a missing portion of the summer assignment. As a reminder, these are the two texts you were given information on a few weeks ago: Campbell Biology, 9e AP* Student Edition (HS Binding) + MasteringBiology with Pearson eText (6-year access) isbn10: 0131375040 isbn13: 9780131375048 AP* Test Prep Workbook for Campbell Biology—Revised for New Curriculum price: $14.97 isbn10: 0321856635 isbn13: 9780321856630 Other Essential Information: You may reach me all summer via my email address (gaetanakalafsky@popejohn.org) or cell (201-213-5727). Please do not hesitate to ask questions. Better to ask up front then to be disappointed in your grades. Do not wait until the last minute to submit your assignments. I have set a time deadline of 11:59 pm for each assignment. This will be 11:59 pm as set by Pope John’s email system. Kalafsky 2 PART 1: Complete Before June 21, 2013 Email the following information to me. Be sure to email it from an account that you WILL check throughout the summer: 1. 2. 3. 4. Name Grade (2013 – 2014) Why did you register to take AP Biology? What are your personal strengths when it comes to learning new material? How do you learn best? 5. What causes you to struggle in a course? 6. What is the most effective way for you to prepare for a test? 7. At this time, what do you plan on majoring in when you get to college? PART 2: Due June 28, 2013 Register for the Mastering Biology site using the access code purchased with your text. 1. 2. 3. 4. 5. 6. 7. 8. Register at www.pearsonschool.com/access Enter the first 6 letters of your code (purchased with your text) Click on Covered Titles to Select Discipline and Title Click on Science Select Campbell, Biology 9e AP* Edition Choose Student Registration Accept - Pearson License Agreement Access Information Create your username & password * Enter your complete access code 9. Account information – complete this section with your name & school information 10. A Confirmation and Summary will be visible on the screen and emailed to you. Register for your specific class within Mastering Biology. 1. Log in to Mastering Biology. 2. Click YES to the question Did you receive a course ID from your instructor. 3. Enter the following Course ID: KALAFSKYAPBIOLOGY20132014 and click Continue. 4. Skip the step that asks you to enter your student ID. 5. You should now be on our specific Mastering Biology Course home page. Kalafsky 3 Learn to use Mastering Biology. 1. On the Assignments tab, complete the Introduction to Mastering Biology assignment. 2. This assignment must be completed prior to June 28, 2012 at 11:59 pm. Note: If you have any difficulty accessing Mastering Biology, I must be notified before June 28, 2013! PART 3: Due August 16, 2013 (by 11:59 pm) AP Biology must cover the topic of Ecology. Since emphasis is placed on understanding concepts, not plain memorizing of facts, you can read the chapters on Ecology as a summer assignment. Ecology is the scientific study of the interactions between organisms and the environment. Because of its great scope, ecology is an enormously complex and also an exciting area of Biology. For each chapter, I have provided additional essats that contain relevant ideas or data on the chapter content. Read the essays that pertain to the chapter, then read the chapter and answer the questions. Question for each chapter should be answered after you have read the article and the chapter. Follow the chapters in order in order to acquire a better understanding of the topics covered in each section. After completing the written questions for each chapter, complete the masteringbiology quiz for the completed chapter. After successful completion of all of the chapters, questions, and mastering biology quizzes, you will be required to complete an free response question (FRQ) related to the topics you have just learned. The FRQ is located immediately in this document, immediately after the chapter questions. In summary: Read Chapters 51 – 56 and the corresponding essays (located in this packet). Complete the attached assignments for Chapters 51 – 56. o Scanned and emailed or mailed to my home by 11:59 pm 8/16/13 Complete the chapter quiz for each chapter (51 – 56). To do this you must log in to the Mastering Biology web site. o To be completed by 8/16/13 at 11:59 pm. Complete the FRQ (found after the chapter questions) o Emailed to me by 11:59 pm 8/16/13 Kalafsky 4 Chapter 51: Animal Behavior Essay - Mating Systems in Sexual Animals 1. How is behavior defined? 2. What is ethology? 3. What is the difference between proximate and ultimate causation? 4. Using red-crowned cranes, provide an example of a proximate causation question and an example of an ultimate causation question? 5. What is a fixed action pattern (FAP)? Provide one example not presented in the text. 6. What is a sign stimulus? Give at least 3 examples of sign stimuli (2 of the examples must not be presented in the text. 7. Nicholas Tinbergen’s work with the stickleback fish is a classic study. Explain what he found. Use the terms fixed action pattern and sign stimulus in your response. 8. Define the both kinesis and taxis and provide one example of each term not presented in the text. 9. Explain what is meant by circadian clock and circadian rhythm. Identify two behaviors, either plant or animal, that demonstrates a circadian rhythm. (You may need to refer to other chapters in the text for examples). 10. Explain two navigational strategies used by birds to migrate. 11. Animals communicate in various ways. Discuss at least three specific examples of animal communication using different organisms. 12. Note Figure 51.4 that shows fruit fly courtship behavior. Identify two other modes of communication used by the fruit fly? 13. Karl von Frisch studied European honeybees. What are the two types of dances that a returning worker bee does, and what information does each dance convey? Use a labeled sketch to describe each dance. 14. What are pheromones? Give three specific types of information that can be transmitted through pheromones. 15. Based on cross-fostering and human twin studies, what are the two factors that contribute significantly to behavior? 16. What is the difference between innate and learned behavior? Give an example of each. 17. What is meant by fitness? How can habituation increase fitness? 18. Describe the process of imprinting, and explain what is meant by sensitive or critical period. 19. Describe the classic study of parental imprinting done by Konrad Lorenz. 20. What occurs in spatial learning? 21. What are two types of associative learning? Which type did Ivan Pavlov use to get a dog to salivate at the sound of a bell? 22. What occurs in operant conditioning? 23. What is cognition? Give three examples of cognition in animal species; include at least one bird behavior. 24. Many bird songs are learned during a critical period. What will happen if a whitecrowned sparrow does not hear the song of its species during this time? 25. What is foraging behavior? Kalafsky 5 26. What is proposed by the optimal foraging theory? Explain it in terms of cost and benefit, and cite two examples from your text. 27. To demonstrate that you understand the principle of optimal foraging, describe a food source that you would not be likely to exploit. 28. Explain each of the following mating systems and provide one example of a species that uses each system. a. Promiscuity b. Monogamy c. Polygamy d. Polygyny e. Polyandry 29. What is sexual selection? (See Chapter 23) 30. There are two types of sexual selection. Explain each of them. a. Intersexual selection b. Intrasexual selection 31. What is agonistic behavior? Give one example of this behavior that is not in your book. 32. What is altruism? 33. Explain the evolutionary advantage to a population of having members who exhibit altruistic behavior. 34. Altruism may reduce the fitness of an individual—for example, by making that individual more obvious to a predator. Explain this behavior using the concept of inclusive fitness. 35. Explain the logic behind the comment that a person would lay down his life for two brothers or eight cousins. 36. Contrast kin selection and reciprocal altruism. Do not simply give a definition. Kalafsky 6 Chapter 52: An Introduction to Ecology and the Biosphere Essay – Global Warming 1. What is ecology? 2. What is a biome? 3. Figure 52.20 shows a climograph for some major biomes in North America. What two abiotic factors shown here are most important in determining the distribution of the biome? 4.Describe each major terrestrial biome as to rainfall, temperature, location, and representative flora and fauna. Biome Rainfall Temperature Location Flora (at least 2) Fauna (at least 2) Tropical Rainforest Desert Savanna Chaparral Temperate grassland Northern coniferous forest/Taiga Temperate broadleaf forest Tundra 5. What is the largest marine biome, and how much of Earth’s surface does it cover? Kalafsky 7 6. As you read this section and study Figure 52.13, you will encounter a number of new terms. Distinguish between each of the following pairs of terms: a. photic/aphotic b.benthic/pelagic c. oligotrophic/eutrophic d.littoral zone/limnetic zone e. zooplankton/phytoplankton f. neritic/abyssal 7. Complete the following chart of the aquatic biomes. Biome Lakes Wetlands Streams and rivers Estuaries Intertidals Ocean Pelagic Description Autotrophs (at least 2) Heterotrophs (at least 2) Negative Human Impact Kalafsky 8 Chapter 53: Population Ecology Essay – Population Ecology, Tales of Nightmare Numbers 1. Draw type I, II, and III survivorship curves on a graph with labeled axes. Explain why the growth rate of species with a type I survivorship curve depends primarily on fertility rates. Explain whythe growth rate of species with a type III survivorship curve is extremely sensitive to changes inadult survivorship. 2. Make a rough sketch of the age distribution in developing versus developed countries, and explain the significance of the differences. 3. Consider 2 rivers: One is spring fed and is constant in water volume and temperature year- round; the other drains a desert landscape andfloods and dries out at unpredictable intervals. Which is more likely to support many species of iteroparous animals? Why? 4. Explain why a constant rate of increase (rmax) for a population produces a growth graph that is J- shaped rather than a straight line. 5. Offer a hypothesis to explain why humans have undergone near-exponential growthfor over 500 years. Why can’t exponential growth continue indefinitely? Give 2 examples of density- dependent factors that influence population growth in natural populations. 6. Where is exponential growth by a plant population more likely- on a newly formed volcanic island or in a mature, undisturbed rain forest? Why? 7. How does the prediction of the exponential model of population growth differ from that of the logistic model? 8. What is carrying capacity? Is it a property of a habitat or of a population? 9. What is time lag? 10. How have humans sidestepped the controls that regulate populations of other organisms? 11. How does the age structure of a population influence its future population growth? 12. Explain why a population that fits the logistic growth model increases more rapidly at intermediate size than at relatively small or large sizes. 13. Identify three density-dependent factors that limit population size, and explain how each exerts negative feedback. Kalafsky 9 Chapter 54: Community Ecology Essay – Community Interactions: No Pigeon is an Island 1. 2. 3. 4. Why are there limits on the food chain length? Compare a dominant species with a keystone species – give an example of each. How do keystone species influence species richness in communities? What are the differences between cryptic coloration, aposematic coloration, and mimicry? Explain the differences. 5. Compare bottom-up and top-down controls on biological communities and their organization. 6. What is disturbance and give an example? 7. To investigate the structure and function of ecosystems, ecologists may construct a microcosm using organisms and materials from the ecosystem. Properly constructed, these model systems should be self-sustaining. If you remove the primary producers from the microcosm, would you predict that your model would continue to be self-sustaining? Explain. 8. If you remove the decomposers and detritivores, would the microcosm be selfsustaining? Explain. 9. Species interactions affect the distribution and abundance of populations. Summarize experimental evidence that population size for snowshoe hares depends on both predation rates by lynx and competition for food among hares. 10. Using your knowledge of ecosystem structure and function, compare the trophic structure of a desert to that of a temperate hardwood forest. Include the relative number (not exact) of organisms and energy availability for the different trophic levels. 11. What is the difference between primary succession and secondary succession? 12. How does the essay, No Pigeon Is An Island, explain the information about community interactions in this chapter? Kalafsky 10 Chapter 55 (Ecosystems and Restoration Ecology) and Chapter 56 (Conservation Biology and Global Change) Essay – Phosphate Pollution, Acid Rain, and the Ozone Hole: Hope for the Ecosystem Recovery Essay – Exotic and Endangered Species 1. Why is the transfer of energy in an ecosystem referred to as energy flow, not energy cycling? 2. How are detritivores essential to sustaining ecosystems? 3. Why is only a small portion of the solar energy that strikes Earth’s atmosphere stored by primary producers? 4. What is the difference between gross primary productivity and net primary productivity? 5. What environmental factors influence rates of primary productivity in terrestrial and aquatic ecosystems? 6. Why is an ecosystem’s net primary production lower than its gross primary production? 7. On a global scale, herbivores consume only about17% of net primary production be terrestrial plants, yet most plant biomass is eventually consumed. Explain. 8. Why is energy lost from a ecosystem at every transfer from one trophic level to the trophic level above it? 9. Marguerite has a vegetable garden in Maine. Eduardo has one in Florida. What are some of the variables that influence primary production in each place? 10. Look around you and name all of the objects, natural or manufactured, that might be contributing to amplification of the greenhouseeffect. 11. Why does deforestation of a watershed increase the concentration of nitrates in streams draining the watershed? 12. Draw a SIMPLE diagram that shows one possible path for an atom or molecule of that chemical from abiotic to biotic reservoirs andback for each of the 4 biogeochemical cycles. 13. How can the addition of excess nutrients to a lake threaten its fish population? 14. In the face of biological magnification of toxins, is it healthier to feed at a lower or higher trophic level? Explain 15. Suppose that herbivores were removed from a temperate deciduous forest ecosystem. Predict what would happen to the rate of nitrogen cycling. Explain the logic behind your prediction. 16. What is Earth’s main reservoir for phosphorus, and why is it recycled at such a slow rate from that reservoir? 17. In 1997, nonnative and invasive Asian swamp eels were collected in Florida for the first time at two sites near Tampa and Miami. These fish are extremely adaptable to a wide range of freshwater habitats, from wetlands to streams and ponds. They are predators that feed on worms, insects, crayfish, frogs, and other fishes, including bluegill and bass. Swamp eels have the ability to gulp air, which allows them to survive in only a few inches of water and to move over land to a Kalafsky 11 nearby body of water. Scientists are tracking their movements and increasing numbers in the Southeast. In one pond, several species of fish have been completely eliminated. Based on your understanding of the pond ecosystem, predict the effect of introducing swamp eels on the following components of the pond. Bluegill: Bass: Pond Life: 18. Using your knowledge of ecosystem structure and function, propose a plan of action for eliminating the swamp eels (question 17) from the pond before they eliminate the other organisms. You cannot use toxins, since the local anglers fish in this pond. 19. In what ways would humans benefit by preserving biodiversity? 20. Describe the 4 main threats to biodiversity and how each one damages diversity. 21. Why does the reduced genetic diversity of small populations make them more vulnerable to extinction? 22. How do naturally occurring organisms provide humans with ecosystem services? 23. What are the consequences of the overexploitation of fish populations? 24. How do extinction rates today compare with the background extinction rate evident in the fossil record? 25. Would a single large nature preserve or several small preserves experience greater edge effects? 26. Why is a population’s effective size (Ne) almost always smaller than its total size (N)? 27. What are the goals of restoration ecology? 28. How do bioremediation and biological augmentation differ? 29. What is meant by the term sustainable development? 30. What are the lessons that can be learned from the essays – use information from the chapter to explain your answer. ECOLOGY FRQ After reading the ecology chapters 51-56 and completing the question listed above, complete the following FRQ. This is a graded essay. Your Answer must be in essay form. Outline form is NOT acceptable. Labeled diagrams may be used to supplement discussion, but in no case will a diagram alone suffice. It is important that you read the question carefully before you begin to write. Your answer MUST be an original work. DO NOT copy from a friend or take information directly from the web. Although your answer may differ from others in the class, it does not mean it is wrong. As long as you answer logically and reasonably provide correct information you will be given credit. Kalafsky 12 These answers do not have to be lengthy. You can answer each part in one paragraph. I am grading on correct information. The food chain can be depicted using arrows between the organisms. FRQ 1. a) Living organisms play an important role in the recycling of many elements within an ecosystem. Discuss how various types of organisms and their biochemical reactions contribute to the recycling of either carbon or nitrogen in an ecosystem. Include in your answer one way in which human activity has an impact on the nutrient cycle you have chosen. b) The survival of organisms depends on regulatory mechanisms at various levels.Explain how the density of a population is regulated. c) Compared with other terrestrial biomes, deserts have extremely low productivity. Discuss how temperature, soil composition, and annual precipitation limit productivity in deserts. d) Describe a four-organism food chain that might characterize a desert community, and identify the trophic level of each organism. Kalafsky 13 ESSAYS FOR AP BIOLOGY SUMMER ASSIGNMENT Essay – Chapter 51 Mating Systems in Sexual Animals One of the most fascinating aspects of human life is how we choose our mates. Animals also choose their mates, sometimes with a great deal of care. Mating systems are important to understand because they reflect the result of natural selection on mate choice, and ultimately on strategies for maximizing individual reproductive success. A mating system describes how males and females pair when choosing a mate. Males and females differ greatly in the investment each makes to reproduce, and may therefore approach mating with differing strategies. To study these differences, scientists observe mating systems and describe how males and females come together. When choosing mates, animals evolve species-typical strategies for maximizing their reproductive success — this results in considerable diversity among animal species in their mating patterns. In this article we first discuss why sexual reproduction exists, and how differences between males and females affect mating systems. We move on to consider the evolution of mate choice, and then we describe the types of mating systems found in animals. The Evolution of Sex Asexually reproducing animals pass on all of their chromosomes, and consequently all copies of each gene, to their offspring. In contrast, due to meiosis, diploid sexually reproducing animals have two copies of each chromosome but only pass one copy of each chromosome on to an egg or sperm cell. This means that a sexually reproducing diploid animal only passes half of its total genes on to its offspring. Despite the cost of losing half of the potential passage of genes to the next generation, sexual reproduction is much more common than asexual reproduction among animals because it provides several evolutionary advantages. The major advantage of sexual reproduction comes from genetic recombination. Genetic recombination allows an organism's offspring to be genetically diverse. Sexual reproduction increases the chances of acquiring favorable mutations and is unlikely to propagate deleterious ones. Genetic diversity within a group of offspring is advantageous as the local environment changes. This idea becomes clear Kalafsky 14 when we examine organisms that can reproduce both sexually and asexually. Aphids, for example, will favor asexual reproduction when their environment is stable. When the environment is going to turn cold, most species of aphids reproduce sexually, because sexual reproduction produces eggs that are freeze tolerant and can diapause during the winter (Simon et al. 2002). Genetic diversity may also lead to evolved defenses against parasites and disease. The mud snail, Potamopyrgus antipodarum, is host to several trematode parasites. Sexual individuals of this species are more common in areas where risk of trematode infection is high. In areas where the risk of infection is low, asexual individuals have displaced sexual ones (King et al. 2009). This suggests that the genetic diversity acquired from sexual reproduction is necessary for this species to defend against parasites, as asexual individuals may not easily survive in areas where parasites are high. Sexual reproduction often involves evolutionary differentiation of males and females. Females typically produce significantly fewer gametes (eggs) than males and invest heavily in each one. On the other hand, males produce many gametes (sperm) and invest little into each one. These strong differences in gamete investment between the sexes leads to reproductive strategies between the sexes that, in some cases, conflict. Females may spend more care than males selecting a mate due to the high cost of their gametes. Figure 1: A male bighorn sheep. Variance in Mating Success and Bateman's Principle A key element of the study of mating systems is understanding how many mates an animal has in its lifetime. Bateman's principle helps to make predictions about mating success and number of mates. Bateman's principle postulates that variance among females in mating success is low, whereas variance among males in mating success is high. This stems from the fact that one mating in females should be enough to fertilize all their eggs whereas in males reproductive success is based on the number of times they have mated. In other words, nearly all females in a population mate and have offspring, Kalafsky 15 but relatively few males mate successfully (Figure 2). Those males that do mate tend to mate with many females-thus a few males have very high reproductive output, but many males have little or no reproductive output (Bateman 1948). This leads to the prediction that sexual selection should act more strongly on males, leading to greater elaboration of behavior and structures used in attracting mates in males than in females. Figure 2: Bateman’s principle. These figures illustrate Bateman’s principle — after one mating, female mating frequency increases and relative fitness remains constant, as the sperm from one mating is adequate to fertilize all the female’s eggs. In males, as mating frequency increases relative fitness also increases proportionally. Criticisms of Bateman's theory focus on the generality of the predictions. Contrary to the predictions of Bateman's principle, there are several possible advantages to female multiple matings. The female cichlid fish Pseudotropheus spiliopterus mates with any male they meet because they have a high risk of getting predated and a small population. This often leads to multiple matings by a single female (Kellogg et al.1998). Mating with any male that is seen ensures that these cichlids have a chance at producing offspring. The female Malawi blue cichlid has a high population but still participates in multiple matings. In this case multiple matings occur to avoid inbreeding and increase genetic diversity among the offspring (Kellogg et al. 1998). Additionally, multiple matings by females may increase the likelihood that they will find a compatible mate, one that is not sterile, or even help prevent infanticide. Kalafsky 16 Female Mate Choice Mate choice is also a key element of mating systems. In most species, females are choosier when picking a mate than males. A significant reason for this is the higher investment females make in each gamete than males. Females may prefer certain males for a variety of reasons, including "good genes", meaning that the male has attributes which predict better survivorship of the offspring, good potential parenting by the male, or possession of resources by the male that will support the offspring during their growth and development. Additionally, in most species, females are more likely to provide parental care. Females that carefully select their mates are at a lower risk of losing their reproductive investment. Males may be under strong selection for certain traits that are favored by females. Most females look at these traits as indicators of their partner's fitness. Selection favors females that choose males that enhance the likelihood of her offspring's success. Males with more elaborate ornamentation, or that are more colorful, can be displaying a good indicator of value as a mate, and may win the chance to mate with a particular female. (Figure 3). Although mating is important, it can be a costly event — females are predicted to be choosier about selecting their mates than males because of risks during mating, such as aggression or disease transmission, which may negatively impact the female's reproductive output. Figure 3: Eyespots from peacock tail feathers. Elaborate ornamentation usually evolves in intrasexual selection and is used in mate choice. Kalafsky 17 Male Mate Choice The importance of male mate choice is controversial. Older theory predicts that male mate choice should be less common in animals. However it plays an important role in many mating systems, and the cost of mating for males may have been underestimated in earlier studies. Male mate choice occurs most often when males are substantially involved in caring for their offspring, or when there is great variation in the quality of the females as mates within a population. If males are choosy about their mate, then over time females may evolve ornamentation or coloration that is subject to sexual selection. Types of Mating Systems Monogamy Social monogamy is the behavioral pairing of a single male with a single female. It is most common in birds and rare in other animals (Figure 4). Theoretically, individuals in monogamous pairs will both contribute to the defense and parental care of offspring. Choosing an inappropriate mate could have a high fitness cost (see the sections above for more on mate choice). Because the costs of poor mate choice in monogamous species can be so high, in some instances organisms engage in strategies of either serial monogamy or extra-pair copulations. Extra-pair copulations are very common in birds (Petrie et al. 1998, Stutchberry 1998). Monogamy reduces the potential for genetic variation among a female's offspring. By mating with more than one male over the course of her lifetime, a female gains higher genetic variation among her offspring. The benefits of monogamy, which are shared parental care and territorial resources, are maintained by having only one mate at a time, or by concealing extra-pair partnerships. Figure 4: Blue-footed boobies. Many bird species, such as these blue-footed boobies are monogamous. Kalafsky 18 Polygyny Polygyny is the association of one male with multiple females. This mating system is found in a few birds and insects, but is most common in mammals. Polygyny is a strategy used by males to increase their reproductive fitness. Resource Defense Polygyny In resource defense polygyny, groups of females are attracted to a resource — males then compete for territorial possession of the resource, and, by extension, mating priority with females at the resource (Beletsky 1994). Thus, individual males form territories centered on resources needed for successful mating (McCracken 1981). Harems Another common type of polygyny is membership in a harem, a defended group of females associated with one male. Females may initially associate in a harem for group defense, or they may be herded together by a male. Males compete for control of the groups. Harems typically exhibit a dominance hierarchy among the females in the group. Leks A lek is an aggregation of males that are each seeking to attract a mate. Within a lek, males typically perform sexual displays. Unlike most other mating systems, leks are not associated with resources. Aggregations of males may be near particularly attractive females or in areas where females are likely to travel (Lank et al. 1995, Aspbury & Gibson 2004). It is thought that males form leks because they attract more females than do isolated males. Attracting more females is a strategy used by males to help increase their reproductive success. Polyandry Polyandry is a group with one female and many males. Polyandry is a reproductive strategy that helps a female ensure reproductive success by providing her with multiple mating options. Resource Defense Polyandry In the Spotted Sandpiper, females control resources, which in turn controls male mating associations (Oring et al. 1994). Cooperative Polyandry The Galapagos hawk exhibits cooperative polyandry. In this case all males in the group copulate with the female and all participate in brood provisioning (Fabborg et al. 1995). Kalafsky 19 Polygynandry Some mating systems have looser male-female bonds within groups. In polygynandrous groups, multiple females and males mate with each other, and males may care for the broods of several females. Chimpanzees and bonobos rely on this strategy — it allows groups of males and females to live together and spend less time being concerned with mate competition. Polygynandry may be advantageous from the female's perspective because it causes paternity confusion, which decreases infanticide and allows her to have multiple males care for her brood (Hrdy 1981, 2000). Promiscuity In promiscuity there are no pair bonds, and males and females, although sometimes choosy, often seem to mate randomly. As it is typically more advantageous for one or both sexes to pick their mate, promiscuity may occur in species for which the environment is unpredictable (Birkhead 2000, Burton 2002). Sperm Competition Although sperm competition is not a type of mating system per se, it is a form of malemale competition that plays an important role in mating systems. If more than one male mates with a female in a short time period, competition can occur after the males have released their sperm (Fisher & Hoekstra 2010). In other words, once a male has released sperm, its sperm must be the first to reach an egg. This is often apparent in animals that use external fertilization. In aquatic animals that release their gametes into the water, animals that release the largest amount of sperm, and sperm that are highly capable of swimming, are likely to produce the most offspring (Stoltz & Neff 2006). Animals with internal fertilization also experience sperm competition. Several mechanisms have evolved to facilitate a male's reproductive success with females that have multiple mates. For example, in one species of damselfly, males physically remove any sperm present from the female before it mates (Waage 1979). Sperm competition adds to the difficulty of obtaining a successful reproductive event by males. Conclusions To transfer their genes to the next generation successfully, animals need to choose a suitable mate. Failure to do so leads to low or no reproductive success — that is, poor fitness. But reproductive success can also hinge on the number of mates, and on social interactions that extend beyond mating. By classifying social interactions, scientists have been able to identify different types of mating systems, such as monogamy and polygyny. The mating systems described in this article represent a variety of strategies to achieve reproductive success. The diversity of mating systems in animals is a fascinating example of the incredible variety of solutions that a complex evolutionary problem can yield. Essay – Chapter 52 Global Warming Kalafsky 20 As the general public and scientists alike become more aware of the suggested patterns and problems of global climate change many begin to realize that not just humans will be affected in the future. We are but one of many species that will be forced to adapt to changing conditions. Those who are unable or fail to adapt will be eliminated by others who can. It is important to understand that regional changes are much more accurate and useful to examine rather than trends on a global scale. Also, when global trends are measurable the changes have already affected the regional elements in huge ways; there exists no reaction time. It is also not realistic to apply these regional changes to all species within its limits. Different species will be affected differently. Consequently, we have chosen birds to narrow our study on how climate change affects species distribution. Birds are excellent indicators of change in spatial distribution and changes have been well documented for birds. We first examined the different problems and issues surrounding global climate change. We then discussed the different theories seen behind the alteration of species distribution due to climate change, and why this is important. We then looked at how different species distributions have altered in general. We concentrated on the change in distribution of birds (as they are a good example of species indicators) including factors as abundance and location, testing if they have changed within the recent past and whether or not these results can be attributed to climate changes. In recent history the world climate has seen significant changes. These changes are not necessarily uncommon as compared with changes of the past, but as far as we can tell the rate and nature of the changes appear to be abnormal. Global climate change includes the theory of global warming or a rise of temperature due to the increase of greenhouse gases in the atmosphere. Scientists have suggested that the earth’s average temperature has increased by as much as 0.6°C (Walther et al., 2002). The estimates on this figure vary and it is projected to rise another 1-3.5°C in the next century (Hughes, 2000). While this might not seem like much, the effects of a small temperature increase can be impressive. The theory behind global warming suggests that increased human activity is putting an abnormally high amount of greenhouse gases (carbon dioxide, water vapor, methane, and nitrous oxide) in to the atmosphere. This can happen through the use of fossil fuels for example, which produce carbon dioxide and methane (Montgomery, 2000). A greenhouse blanket is formed from these gases, that warms and allows the earth to sustain life, and it is growing ever thicker. When this happens, the greenhouse gases trap longerwavelength infrared rays that would otherwise be able to be radiated out into space (Montgomery, 2000). This in turn leads to containing heat within the region between this blanket and the earth, thus elevating the earth’s temperature (NZCCO). This process is known as global warming and leads to potential climate changes. Other changes in precipitation and wind will likely be associated with the temperature changes. “Associated changes in wind-flow patterns and amounts and distribution of precipitation will cause differential impacts in different areas, not all of which will be equally resilient” (Montgomery, 2000). All these changes in climate alter many other aspects of the natural environment; everything from alterations in ice sheets and glaciers to trended movements of biomes and species distribution. It can be seen that global warming is a serious phenomena, potentially affecting sea levels, glaciation, agriculture, ocean cycle, human health, species spatial distribution, and much more. Kalafsky 21 Global warming is a complicated process with many causes and effects that are difficult to predict or control. At the current time it is the general consensus of the scientific community that global warming and the increase in temperature are largely due to human action and influence. There exist two types of green house effects; the natural greenhouse effect and the enhanced greenhouse effect. The natural greenhouse effect prevents the earth from being frigid and inhospitable. It’s because of these gases that life on this planet exist, “The natural greenhouse effect causes the mean temperature of the earth’s surface to be about 33 degrees C warmer than it would be if natural greenhouse gases were not present (NASA 2).” The enhanced greenhouse effect is the unnatural effect that was brought on by humans. “It’s the possible raising of the mean temperature of the Earth’s surface above that occurring due to the natural greenhouse effect (2)”. The greenhouse gases that are the culprits in the enhanced greenhouse effect are mainly water vapor, ozone, carbon dioxide, methane, nitrous oxide, and chlorofluorocarbons (CFC’s). Carbon dioxide along with the other gases have been increasing in the atmosphere over the past couple centuries, “There has been about a 25 % increase in carbon dioxide in the atmosphere from 270 or 280 ppm 250 years ago, to approximately 360 ppm today (3).” These records and recordings of the abundance of carbon fluctuate depending on the hemisphere you’re in, the season it is, and at what time of day it is. Interestingly enough the two most abundant gases in the atmosphere, nitrogen and oxygen, have no effect on the greenhouse warming (unless when combined in the compound of nitrous oxide). As greenhouse warming continues, there are some processes that seem to counter the downward spiral of global warming. One example of these processes is in relation to the carbon cycle. It has been noted that in the global plant community many plants seem to be taking up more carbon dioxide and storing it in either biomass or soil. It is doing this in association with the increased carbon dioxide that is present in the atmosphere. Plants are therefore a sink for the excess carbon dioxide. This process has different implications. “Air temperatures over the land have increased, resulting in a lengthened growing season in the northern and mid-latitudes; gradual and slight warming seems to have favored photosynthesis over respiration-decomposition with far-reaching effects on the global carbon balance” (NASA FACTS 2). Other processes such as this sink for carbon dioxide are being studied as to how much of an impact they can have to counter global warming and their limits on their effectiveness. An understanding of the current climatic regions and resulting biomes is important for studying species distributions and changes of distributions. In a traditional system of physical geography, the world can be broken down into eleven terrestrial biomes based on vegetation classes. These include equatorial and tropical rain forest, tropical seasonal forest and scrub, tropical savanna, midlatitude broadleaf and mixed forest, needleleaf forest and montane forest, temperate rain forest, mediterranean shrubland, midlatitude grasslands, warm desert and semidesert, cold desert and semidesert, and arctic and alpine tundra (Christopherson, 2003). These biomes are characterized by the vegetation, soil, climate including precipitation range, temperature, and water balance. Latitude plays a big part in determining these biomes as you can tell by some of their names. The most intense or distinctive biomes are the tropical rain forest, the deserts, and the alpine and arctic. The tropical rain forest has “consistent year-round daylength (12 hours), high insolation, average annual temperatures around 25°C (77°F), and plentiful moisture, plant Kalafsky 22 and animal populations have responded with the most diverse expressions of life on the planet” (Christopherson, 2003). Average annual rainfall in a rain forest is at least 80 inches and some can have as much as 200 inches per year (Weigle, 1999). There is no dry season. Tropical rain forests can be found in four main regions: Central and South America, West and Central Africa, South and Southeast Asia, and Australia (Weigel, 1999). Although the rain forest soils support a vast array of plant life, they are lacking in nutrients because the plants consume the nutrients so quickly (Weigle, 1999). Common rain forests plants include bamboo, hibiscus shrubs, orchids, African violets, ferns, and Spanish mosses (Weigle, 1999). Some of the common rain forest trees are black ebony, cinchona, mahogany, and mango (Weigle, 1999). Invertebrates are vastly abundant in the rain forest and include such organisms as termites, army ants, orchid bees, birdwing butterflies, postman butterflies, and hunting wasps (Weigle, 1999). Reptiles and amphibians such as frogs, toads, salamanders, snakes, and chameleons are commonly found here (Weigle, 1999). Mammals of the rain forest include monkeys, shrews, bats, sloths, gorillas, and jaguars (Weigle, 1999). The rain forest has larger bird populations than any other biome and includes some of the following: hummingbirds, birds of paradise, jacamars, eagles, parrots, and junglefowl (Weigle, 1999). The desert regions are created by descending, drying, and stable air of high-pressure systems from 8 to 12 months of the year. On average, deserts receive less than 10 inches of rainfall a year (Weigle, 1999). Here evaporation always exceeds precipitation (Weigle, 1999). Temperatures may reach between 105-110°F during the day in hot deserts (Weigle, 1999). However, night temperatures can fall to 50°F (Weigle, 1999). Desert soils are coarse, light colored and high in mineral content (Weigle, 1999). Some deserts have little soil but rather pebbly rock or desert pavement (Weigle, 1999). Because of these features, the organisms that live in the desert must be adapted to these conditions. Consequently for example, many plants have evolved a waxy coating on them to prevent water loss. Although deserts do not support large numbers of plants and other organisms, they can support a wide range of organisms (Weigle, 1999). “Cold deserts have hot summers and cold winters (Weigle, 1999)”. Cold deserts can be found in Kazakhstan, Uzbekistan, China, Mongolia, and Utah (Weigle, 1999). They can receive precipitation in the form of snow (Weigle, 1999). The alpine or arctic regions can have conditions of almost continuous daylight or continuous darkness for up to two months due to its high latitude. This phenomenon has to do with the earth’s tilt and overall orbit and the tundra’s location above the Artic Circle. “Arctic tundra is found across northern Alaska, Canada, and Siberia (www.cotf.edu)”. Winters here are long and cold while summers are cool and brief. “Intensely cold continental polar air masses and stable high-pressure anticyclones govern tundra winters. A growing season of sorts lasts only 60-80 days, and even then frosts can occur at any time (Christopherson, 2003)”. There are low levels of precipitation, similar to that of the desert, and dry winds are common (www.cotf.edu). “Vegetation is fragile in this flat, treeless world; soils are poorly developed periglacial surfaces, which are underlain by permafrost (Christopherson, 2003)”. The permafrost cannot be penetrated by roots or water. There are not many animals that live year-round in the tundra. Birds and mammals such as the artic wolf, brown bear, and muskox, come to live here in the summer (www.cotf.edu). Mosses, lichens, and grasses can grow in the artic tundra (www.cotf.edu). Kalafsky 23 The rest of the biomes are mild or moderate in comparison to those already described, with conditions of temperature and precipitation and consequently organisms ranging on a spectrum within these extremes. Grasslands are characterized by having the dominant plants that are grasses rather than trees or shrubs (Weigle, 1999). Grasslands cover up to 25-30% of the earth and are found primarily in the interior of the continents (Weigle, 1999). They are usually windy and dry for part of the year and are found primarily on flat or gently rolling hills (Weigle, 1999). Grasslands are considered transition zones between deserts and forests (Weigle, 1999). The midlatitude broadleaf, or deciduous forest, can be found in the eastern U.S., central and western Europe, Russia, Japan, and China (Weigle, 1999). Common types of trees here include beech, maple, oak, hickory, ash, and birch (Weigle, 1999). The soils in a deciduous forest are rich (Weigle, 1999). Needleleaf, or coniferous forests often have acidic soils due to the high acid content of the needles (Weigle, 1999). “More than 50 percent of the world’s coniferous forests are found in Asia, primarily in Siberia, China, Korea, and Japan, and on the slopes of the Himalaya and Hindu Kush Mountains (Weigle, 1999)”. They can also be found in Europe, North America, and some in South America (Weigle, 1999). Scientists have begun to study how the changes in climate are affecting a wide range of species. There have been a wide variety of groups including plants and animals, vertebrates and invertebrates, terrestrial and aquatic, and tropical and polar species that have been monitored. They have looked at everything from physiology, phenology, distribution, adaptation, community, and ecosystem structure. Climate plays an important role when it comes to the distribution of organisms as can be seen by studying the biomes. Significant shifts in distribution have been seen before as climate shifts have occurred. Many organisms are able to shift with the changing climate and adapt to the modification, while others cannot. Many of those who can’t are disabled by three main causes; when there are reductions in species diversity through reductions in habitat size, when warming exceeds the migrational capabilities of a species, and when there are losses of habitat during progressive shifts of climatic conditions (GWTBD 4). The basic theory behind the decline in patch area and associated species loss is associated with the theory of island biogeography. “Biogeography is the study of the distribution of plants and animals, the diverse spatial patterns they create, and the physical and biological processes, past and present, that produce Earth’s species richness” (Christopherson, 2003). When further expanded to habitat destruction and disturbance of islands or isolated areas, it is predicted that species diversity will decrease with decreasing island size (5). This theory is thought to hold true similarly in reference to a patch of forest or even in the ocean. When discussing the dilemma of when warming exceeds migrational capabilities of species there is much that we cannot understand or calculate. No one, at this point, is entirely sure at what rate species will be able to migrate under such stresses. This makes it difficult to predict a likely outcome if these events were to happen. Instead of actually predicting how fast organisms will move, we can predict how fast organisms will have to move in order to keep up. With the application of already established models able to predict current and future distributions of major vegetation types, we can predict how fast biomes will shift. With this accomplished, we could also get a general understanding of how fast the species found within each of the biomes would also have to move. The Kalafsky 24 species’ survival depends on their ability to remain in their shifted biome; it is unlikely that they would survive in another biome. Fortunately, there are transition zones between the biomes that allow for some leeway. The third event that species may have to deal with is the loss of habitats during progressive shifts of climatic conditions. With this issue, the main element restricting species movements is human induced. When biomes shift they could potentially shift into areas that are fragmented by human variations of land use. For example, land that is currently used for agriculture or urban areas would be uninhabitable for most species. This fragmentation effect of these areas could potentially result in species loss (GWTBD). “The pivotal importance of patchiness in the ecology of individuals, populations, and communities is now widely recognized (Perrins et al 1991)”. Some distinct habitats in particular have been greatly destructed. These include estuaries, ancient woodlands, and moorlands (Perrins et al, 1991). For conservation practices, we must be concerned with both the size of patches and their separation (Perrins et al, 1991). “The importance of distance between patches is emphasized by our analysis of the cost of dispersal or migration between patches (Perrins et al, 1991)”. The relative percentages of human disturbance by continent are displayed in the following figure(Chapin et al, 2001). According to the following figure, biodiversity continues to decrease as the negative factors including mostly all human induced changes (directly or indirectly) overpower the positive factors contributing to biodiversity (Chapin et. al, 2001). Kalafsky 25 Specific examples of a variety of organisms are useful in observing trends of climate change on species distributions. It is often predicted that organisms will move poleward to compensate for the overall warming of temperature. Plants are interesting organisms to monitor because they can’t physically get up and relocate as animals can, yet changes in distributions have been observed. Plants require a certain amount of warmth to complete their life cycle (Chapin et al, 2001)”. This “amount of warmth” can be measured by a certain number of days above a certain temperature for example (Chapin et al, 2001)”. Dispersal patterns and survival in the new environments are starting to be studied. For example, tree species in the eastern U.S. have been documented to have moved poleward (Iverson & Prasad, 2002). Unfortunately, it has been found that some plant species are unable to establish populations in new areas when faced with climate change and habitat fragmentation (Primack & Miao, 1992). Therefore, some of these plant species are experiencing population declines or even extinctions. Lichens have also been monitored and it seems that species that ordinarily have a subtropical distribution are invading more poleward areas (van Herk et al. 2002). This poleward movement seems to make sense in order to compensate for the temperature increase. With a latitudinal movement toward the poles, there would likely be a few degrees decrease in temperature for the organism. Kalafsky 26 The effects on animals can be studied more specifically in either terrestrial or aquatic settings. The intertidal zone is also a great region to study. This zone is the transition between the aquatic and terrestrial zone. These organisms are already subject to conditions of great stress. They are subject to periods of desiccation and inundation, and they have to deal with the salinity of the ocean as well. It has been proposed that the animals in this region will be most affected by the effects of global warming. Stillman has studied the effects of temperature changes in crabs in the intertidal zone (2002). In addition to all the other conditions that these crabs must be adapted to, with temperature change added in, there will likely be negative effects on the population. Insects are also an important group of animals to consider. Changes in insect distributions can lead to other human implications through the spread of diseases. For example, mosquitoes have been reported at higher altitudes in Latin America and Africa, areas which already have high instances of malaria (Hughes, 2000). With their population shift to higher latitudes, people at these latitudes may be increasingly affected by the diseases such as malaria that they carry. Lastly, corals are a very important example to look at for the effects of climate change. They are already extremely sensitive organisms who are affected by even slight natural changes in salinity or temperature. There have been many studies done to see the effects of global warming on corals and the evidence suggests mass extinctions of certain species (Hughes, 2000). There have been massive bleaching effects observed in corals already. This bleaching happens when the zooxanthellae, the symbiotic algae critical for the corals’ survival, leave the corals. Although the mechanisms of bleaching are poorly understood, it is believed to be linked to temperature, salinity, toxicity, or light, or a combination of these varied effects (Davidson, 1998). “Corals are, in a sense, the highwire artists of the sea, where only a slight misstep spells doom” (Davidson, 1998). We have narrowed down our study from these categories and have chosen to primarily look at the effects of climate change bird species distribution. “It has been argued that if bird communities are protected, then many other communities will be protected, and that evaluation of bird communities can be used as an indicator of the quality and Kalafsky 27 conservation interest of the habitats (Spellerber, 1992)”. Birds are also becoming increasingly well documented due to the international interest and cooperation of many different countries and organizations (Spellerberg, 1992). Although population statistics are for birds are difficult due to their mobile lifestyles (Spellerberg, 1992), birds are excellent indicators of climate change because effects can be seen quickly. They have the ability to relocate to more favorable or suitable conditions. “Bird species richness has been related to vegetation height diversity (Chapin et al, 2001)”. Consequently, one would expect bird populations to shift as vegetation patterns are shifting. Some studies have found that bird distributions are most affected by climate and altitude (Storch et al, 2003). The climate factors have been studied by many and have found that not only do bird distributions vary based on temperature but also largely on precipitation as well (Githaiga-Mwicigi, 2002). The following graphs are from a study done on birds in Mexico and show that there is a correlation between bird distribution and temperature and precipitation (Gomez da Silva & Medellin, 2002). The first graph had a p-value of <0.001 which indicates that temperature did have a significant difference on bird species abundance (Gomez da Silva & Medellin, 2002). The second graph had a p-value of 0.006, again indicating a significant effect of precipitation on bird species abundance (Gomez da Silva & Medellin, 2002). Birds have been subjected to changes due to habitat degradation along with these indirect effects from humans such as climate change and other disturbances. When these factors are coupled with limitations based on specialization, certain species are more influenced than others (Julliard et al 2003). It is suggested that island or mountaintop species will be particularly vulnerable to climate changes (Newton, 1998). As a general rule, if a habitat is reduced in area by 50%, about 10% of the species will be lost, and if the habitat area is reduced by 90%, about 50% of the species will be lost (Newton, 1998). Organisms in tropical forests are in particular danger due to their high rates of habitat destruction Kalafsky 28 (Newton, 1998). The importance of land area to characteristics such as species richness can be seen in the following graph (MacDonald & Kirkpatrick, 2003). Some of the island species that are now considered threatened are following the patterns of those that have gone extinct in the last 400 years. Included in these are tube-nosed petrels (Procellariidae), frigate birds (Fregatidae), pigeons (Columbidae), honey-eaters (Meliphagidae), honey-creepers (Drepanididae), and wattlebirds (Callaeidae) (Newton, 1998). The Pacific Ocean alone is home to 110 of these threatened island species (Newton, 1998). The following figure breaks down the threatened and extinct birds by island regions (Newton, 1998). Kalafsky 29 Contrastingly, some species of birds have been affected by the human suppression of natural disturbances. For example, bird ecosystems are often regulated by events such as fires or droughts. As humans subdue these natural disturbances, the ecosystems are bound to be impacted (Brawn, et al. 2001). These natural disturbances are necessary for regulating the environmental conditions of an area. For example, fires result in the limiting of trees in grasslands as well as providing ashes which make fertile soil after recovery. As natural disturbances help regulate and further characterize vegetation in biomes, land-use is also studied in relation to species distribution as it directly impacts vegetation. Venier et al. found that indeed land-use patterns do influence the distribution of certain species of birds (2004). “With the increasing population pressures and the development of modern agriculture, the sustainable use of biodiversity has lost its role in these systems. This trend has rapidly led to the destruction of local and regional biodiversity in agricultural systems and marginalized natural vegetation and wildlife as a natural resource (Chapin et al, 2001)”. Land use change was found to be the driver with the biggest impact on biodiversity according to biome model scenarios, followed by climate change as the next most important (Chapin et al, 2001). This has to do primarily with the greater impacts at higher latitudes. The following table illustrates their findings on the drivers of biodiversity (Chapin et al, 2001). Kalafsky 30 A negative consequence of the suppression of natural disturbances is that they are likely to be more extreme and abundant as a side-effect. For example, with the suppression of natural fires, land is likely to be drier and thus further promote larger fires in the future. The prediction for increase in fires in the future is displayed in the following image (Chapin et al, 2001). The migratory patterns of birds can be tracked and their nests serve as important studies on their hatching patterns. It has been suspected that breeding ranges would move upward Kalafsky 31 in latitude or elevation due to the increase in temperatures. However, from some of the studies so far, this theory has not yet been supported (Archaux, 2004). Bird experiments have been well documented worldwide and thus we feel that they are an excellent group to study in order to see how climate change can directly affect certain species’ distributions. From one of the studies that we found, from the Rocky Mountain Bird Observatory, noticed what they found to be unusual bird patterns for the 2001 year. From their bird banding study, they found high numbers of the following species: Western Tanagers, Bullock’s Orioles, Yellow-rumped Warblers, and Western Wood Peewee (Boulder County Nature Association, 2001). Some of the species that they found low numbers or entirely missing include Northern Waterthrush and Blue Grosbeak (Boulder County Nature Association, 2001). We chose five of the species from this study that we knew to be migratory birds and compared their abundances from their data from 19912001. The species that we chose were the American Goldfinch, American Tree Sparrow, Brown-headed Cowbird, Gray Catbird, and the House Wren. The American Goldfinch is found in southern Canada and southern U.S. It migrates primarily in flocks during the day (Bird Index). The American Tree Sparrow can be found in the summer in Alaska and northern Canada and in the winter in southern Canada and the central U.S. (Bird Index). The Brown-headed Cowbird is found primarily in North America, specifically in the U.S. and Mexico and is known to migrate shorter distances (Chipper Woods Bird Observatory). The Gray Catbird is found in southern Canada and eastern and central U.S.(Bird Index). It American spends its winters in Panama and the West Indies. The Gray Catbird migrates at night (Bird Index). The House Wren is commonly found from central Canada to southern South America and in the summer specifically in the U.S. and Canada (Bird Index). Kalafsky 32 Fig 1. - contrasts bird abundance over time of five different migratory species common in North America Kalafsky 33 Although there seems to be great variability, there are some patterns worth noting. This graph was taken from data collected semi-annually in the spring and fall between the years of 1991-2001. The years of 1991-1995 were grouped and graphed together and consequently most species had high numbers for the first 2 data points (corresponding to 1991-1995 spring and 1991-1995 fall, despite the label). Every increment after that goes up by a half-year accordingly. The March and September labels were arbitrarily picked to represent spring and fall respectively. Three of the species, the American Goldfinch, the Gray Catbird, and the Brown-headed Cowbird had low numbers in the fall (September) of 2000. We are attributing this to the wildfires in Colorado at this time. This makes sense due to the location of the observatory (Rocky Mountain Observatory). Another low time was in the fall of 1997 for all species. A number of bird species in this area show a slight decrease in population, but as to why this slight decrease occureed the evidence is inconclusive. Wildfires could potentially account for the drop off in 2000, but as for the less dramatic fluctuations in population there could be many other compounding factors influencing these events. The overall slight decline in bird populations could be representative of events where birds are shifting out of their common habitats. One would think that other populations of birds would move into the regions that were being moved out of, but in the mountains of Colorado, where this study was conducted, the actual presence of the mountains may be acting as geographic barriers impeding the other species abilities to move into the given region. It is important to understand how species numbers are estimated in order to know what management approaches should be taken. One model for the population studies of birds involves the following equation in their model: CV=_ni/Ni (Spellerberg, 1992). Here, CV is the index, ni is the number of pairs of the ith species at a sit and Ni is the population size of the ith species (Spellerberg, 1992). This study was done primarily in Western Europe. Another more common one is the Shannon-Weiner indes of species diversity. This model uses the equation H’= - _ pi (logPi) where Pi is the proportion of individuals of species i in the sample (Spellerberg, 1992). These two models along with many others have created useful data for the assessment of particular habitats. From these, the IUCN and the IWRB (International Waterfowl Research Bureau) have identified internationally important wetlands of Europe and North Africa (Spellerberg, 1992). Some of the criteria used to identify an important wetland for the populations include: regularly supporting 1% of the biogeographical population of one species, supporting an appreciable number of an endangered species of plant or animal, or playing a major role in that region for scientific or economic importance (Spellerberg, 1992). We conclude that species spatial distributions are directly effected by global warming and subsequently climate change. In general terms it has been stated by the scientific community that the distribution of species have been moving in a poleward trend. Within the realm of our study we found no conclusive evidence to prove or disprove this statement. The evidence that we did find and cited leads us to the conclusion that the distribution of species is infact being altered by climatic change, but we were unable to determine exactly what that change was. This project focused on bird species (as we found they were ideal indicators of species shifts due to the fact that their patterns of movement are already larger and more immediate than other organisms. This and the fact that bird movements and migrations are well documented are the reason we chose to focus our study on birds). Evidence found specifically from birds shows that there is a Kalafsky 34 correlation between bird population characteristics and alterations in climatic factors such as temperature and precipitation. The change in population characteristics shows that some sort of shift or generally trended movement is occurring. To contribute to the sustainable future which we all would like to see, we must work on reducing our ecological footprint on the earth. An ecological footprint is “A measure of human pressures on the natural environment from the consumption of renewable resources and the production of pollution (Knox & Marston, 2004)”. This means trying to conserve as many of the species of birds, and other organisms alike, for future generations to come. The idea of sustainable development is “A vision of development that seeks a balance among considerations of economic growth, environmental impacts, and social equity (Knox & Marston, 2004)”. This should be a model for all nations and organizations alike; to create a future in which a balance exists between the physical and human worlds. Management and conservation practices can make a difference. For example, Newton illustrates examples of bird species (with their associated distribution ranges) that have seen significant results in increasing population sizes due to management practices since 1950 (1998). Among these include the Whooping Crane, California Condor, Hawaiian Crow, and Crested Ibis, just to name a few (Newton, 1998). Newton attributes the success of these species to protection of the existing populations and ensuring suitable habitats (1998). Most remarkable was the case of the Mauritius Kestrel. This species in 1974 only had four individuals remaining (Newton, 1998). Through the success of a captive breeding and release program, the number of individuals rose to 200+ by 1993 (Newton, 1998). From these examples, one can see that there is a chance to save the species which are most affected by human disturbance and climate change. A good background and understanding of species distributions is crucial for management and conservation techniques. With greater public awareness and concern, we could alleviate some of the problems that species are facing in response to global climate change; however, action must be taken on the regional or local level to be successful. As human population continues to increase, there will consequently be increasing impacts on species worldwide. Habitat destruction and climate change will force species into regions or situations that they will be forced to adapt to. Failure to adapt will lead to declining populations and could result in a great loss of biodiversity. It is important to acknowledge the impact that humans are having on species and also to work towards reducing our "ecological footprint". This means countering destructive practices that humans have engaged in for centuries. This is key for ensuring a sustainable future for coming generations. Kalafsky 35 Essay – Chapter 53 Tales of Nightmare Numbers Across from Sausalito, California, the steep flanks of Angel Island rise form the waters of San Francisco Bay. The island, set aside as a game reserve, escaped urban development. It did not escape from the descendants of a few deer that well-meaning nature lovers shipped over in the early 1900s. With no natural predators to keep themin check, the few deer became many-far too many for the limited food supply of their isolated habitat. Yet the island attracted a steady stream of picnickers from the mainland. They felt sorry for the malnourished animals and made sure to load the picnic baskets with extra food for them. The visitors imported so much food that scrawny deer kept on living and reproducing. In time, the herd nibbled away the native grasses, theroots of which had helped slow soil erosion on the steep hillsides. Hungry deer chewed off all the new leaves of seedlings; they killed small trees by stripping the bark and its phloem. The herd was destroying the environment. In desperation, game managers proposed using a few skilled hunters to thin the herd. They were strongly denounced as being cruel. They proposed importing a few coyotes to the island to thin the herd naturally. Animal rights advocates opposed that solution, also. As a compromise, about 200 of the 300+ deer were captured, loaded onto a boat, and shipped to suitable mainland habitats. A number of them received collars with radio transmitters so that game managers could track them after the release. In less than 60 days, dogs, coyotes, bobcats, hunters, and speeding cars and trucks had killed off most of them. In the end, relocating each surviving deer had cost taxpayers almost 3,000 dollars. Kalafsky 36 The State of California refused to do it again. And no one else, anywhere, volunteered to pick up future tabs. It is not difficult to define the boundaries of Angel Island or track its inhabitants, so it is easy to draw a lesson from this tale: A population’s growth depends on the resources of its environment. And attempts to “beat nature” by altering the sometimes cruel outcome of limited resources only postpone the inevitable. Does the same lesson apply to other populations, inother places? Yes, it does, as the next tale makes clear. When 1999 drew to a close, there were over 6 billion people on Earth. About 2 billion already live in poverty. Each year 40 million more join the ranks of the starving. Next to China, India is the most populous country, with more than a billion inhabitants. By 2010 there may be 182 million more. Forty percent of those people live in rat-infested shantytowns, without enough food or fresh water. They are forced to wash clothes and dishes in open sewers. Land available to raise their food shrinks by 365 acres a day, on average. Why? Irrigated soil becomes too salty when it drains poorly, and there is not enough water to flush away the salts. Can wealthier, less densely populated nations help?After all, they use most of the world’s resources. Maybe they should learn to get by more efficiently, on less. For example, people might limit their meals to cereal grains and water; give up their private cars, living quarters, air conditioners, televisions, and dishwashers; stop taking vacations, and stop laundering so much; close all the malls, restaurants, and theatres at night; and so on. Maybe wealthier nations also should donate more surplus food than they already donate to less fortunate ones. Then again, would huge donations help, or would they encourage dependency and spur more increases in population size? And what if surpluses run out? In is a monumental dilemma. At one extreme, the redistribution of resources on a global scale would allow the greatest number of people to survive, but at the lowest comfort level. At the other extreme, foreign aid rationed only to nations that restrict population growth would allow fewer individuals to be born, but the quality of life would be greater. Currently, the foreign aid program of the United States is based on two premises: (1) that individuals of every nation have an irrevocable right to bear children, even if unrestricted reproduction ruins the environment that must sustain them; and (2) that because human life is precious above all else, the wealthiest nations have an absolute moral obligation to save lives everywhere. Regardless of the positions that nations take on this issue, ultimately they must come to terms with this fact: Certain principles govern the growth and sustainability of populations over time. These principles are the bedrock of ecology – the systematic study of how organisms interact with one another and with their physical and chemical environment. Ecological interactions start within and between populations, and they extend on through communities, ecosystems, and the biosphere. Kalafsky 37 Essay – Chapter 54 NO PIGEON IS AN ISLAND Flying through the rain forests of New Guinea is an extraordinary pigeon with cobalt blue feathers and lacy plumes on its head. It is about as big as a turkey, and it flaps so slowly and noisily that its flight sounds like an idling truck. As is true of eight species of smaller pigeons living in the same forest, it perches on branches to eat fruit. How is it possible that nine species of large and small fruit-eating pigeons live in the space of the same forest? Wouldn’t you think that competition for food would leave one the winner? In fact, in that rain forest, every species lives, grows, and reproduces in a characteristic way, as defined by its relationships with other organisms and with the surroundings. Big pigeons perch on the sturdiest branches when they feed, and they eat big fruit. Smaller pigeons, with their smaller bills, cannot open big fruit. They eat small fruit hanging from slender branches that are not sturdy enough to support the weight of a turkey-sized pigeon. The species of trees in the forest differ with respect to the diameter of their fruit-bearing branches and the size of their fruit. So they attract different pigeons with different characteristics. Is such ways, the nine species of pigeons partition the fruit supply. And how do individual trees benefit from enticing the pigeons to dine? The seeds inside their fruits have tough coats, which can resist the action of digestive enzymes inside the pigeon gut. During the time it takes for ingested seeds to travel through the gut, the pigeons fly about, so they dispense seed-containing droppings in more than one place. In this way, the pigeons tend to disperse seeds some distance from the parent plant. Later, when seedlings grow, the odds are better that at least some will not have to compete with their parents for sunlight, water, and nutrients. Seeds that drop close to home cannot compete in and significant way with the resource-gathering capacity of the mature trees, which already have extensive, well-developed roots and leafy crowns. Within the same forest, leaf-eating, fruit-munching, and bud-nipping insects interact with other organisms and their surroundings in certain ways. So do nectar-drinking, flower pollinating bats, birds, and insects. And so do great numbers of beetles, worms, and other invertebrates that busily extract energy from remains and wastes of other organisms on the forest floor. By their activities, they cycle nutrients back to the trees. Like humans, then, no pigeon is an island, isolatedfrom the rest of the living world. The nine species of New Guinea pigeons eat fruit of different sizes. They disperse seeds from different sorts of trees. Dispersal influences where new trees will grow and where the decomposers will flourish. Ultimately, tree distribution and decomposition activities influence how the entire forest community is organized. Directly or indirectly, interactions among coexisting populations organize the community to which they belong. Kalafsky 38 Essay – Chapter 55/56 PHOSPHATE POLLUTION, ACID RAIN, AND THE OZONE HOLE: HOPE FOR THE ECOSYSTEM RECOVERY? A central message of chapter 54 is that energy flows and nutrient cycling are undergoing extraordinary changes in ecosystems throughout the world. Although the outcome of these changes is uncertain, it is important to recognize that humans have already identified and acted on several recent changes in the abiotic environment that clearly had negative consequences. The events and responses took place at the local, regional, and global levels. The first example of an effective response to an abiotic change took place at a local level and involved a global nutrient cycle. Like many other elements and molecules, phosphorus cycles through ecosystems. The use of phosphate-based detergents in the industrialized countries led to a large increase in the concentrations of phosphate in lakes and streams, triggering rapid and widespread eutrophication – particularly in shallow lakes that received out-flow from municipal sewage systems. In response, governments in North America and Western Europe encouraged or required the use of phosphate-free detergents, and sewage plants were upgraded to remove more phosphate during treatment. Although phosphate pollution from farm fertilizers remains a serious problem, the crisis conditions of the 1960’s and 1970’s have largely been alleviated. The second example involves changes in the pH of rainwater at a regional level. The problem began with sulfur oxides and nitrogen oxides that are pumped into the atmosphere by coal burning electrical power plants and vehicles that lack catalytic converters. When exposed to sunlight and water vapor, the molecules react to form sulfuric acid (H2SO4) and nitric acid (HNO3). Normal rainwater has a slightly acidic pH, about 5.6. But in areas affected by acid rain, precipitation can have a pH as low as 4.2 or 4.4. During the 1980’s and the 1990’s, biologists documented that forests and lakes in eastern North America and northern Europe were being affected by acid rain. Tree growth slowed in response to the acidification of soils, and lakes became less productive and less diverse. Once biologists had documented the problem, governments instituted stricter controls on the amounts of sulfur oxides and nitrogen oxides that could be emitted from power plants, cars, and trucks. Over the past decade, the intensity of acid rain has diminished and some of the ecosystems that were being affected have begun to recover. A third example involves changes in atmospheric chemistry that were global in scale. Widespread use of the compounds called chlorofluorocarbons (CFCs) in refrigeration and aerosol products resulted in the release of thousands of tons of CFCs into the atmosphere. When CFCs accumulated in the upper atmosphere, they participated in chemical reactions that released chlorine atoms. These chlorine atoms subsequently reacted with ozone (O3) molecules, which also accumulate in the upper atmosphere. In some years, the loss of ozone due to these reactions was so severe that an ozone hole opened over Antarctica. This issue concerned scientists from around the world, because ozone absorbs large amounts of ultraviolet (UV) radiation. When the ozone layer that surrounds Earth thins or is wiped out, an excess of UV radiation can reach Earth’s surface and act as a mutagen and carcinogen. Fortunately, soon after the problem was documented in the Kalafsky 39 early 1990’s, international treaties scheduled and enforced the rapid phasing out of CFC productions and use. Scientists have recently been able to document the first signs that the size and duration of the ozone hole may be moderating. The message of these examples is clear: Effective responses have occurred at the local, regional, and global levels when biologists documented serious problems in ecosystem ecology. It remains to be seen whether the same success can be achieved in response to global warming, nitrate pollution, and other current problems in the abiotic environment. Essay – Chapter 55/56 EXOTIC AND ENDANGERED SPECIES When you hear someone bubbling enthusiastically about an exotic species, you can safely bet the speaker is not an ecologist. This is a name for a resident of an established community that has moved from its home range and successfully taken up residence elsewhere. It makes no difference whether the importation was deliberate or accidental. Unlike most imports, which cannot take hold outside their home range, an exotic species insinuates itself into the new community. Sometimes the additions are harmless and even have beneficial effects. More often, they make native species endangered species, which by definition are extremely vulnerable to extinction. Of all species that are now on rare or endangered lists or have already become extinct, close to 70 percent owe their precarious existence or demise to displacement by exotic species. HELLO VICTORIA, GOOD-BYE CICHLIDS. Finding better ways to manage our food supplies is essential, given the astounding growth rate of the human population. Such efforts are well intentioned, but they can have disastrous consequences when ecological principles are not taken into account. For example, several years ago, someone thought it would be a great idea to introduce the Nile perch into Lake Victoria in East Africa. People had been using simple, traditional methods of fishing there for thousands of years. Now they were taking too many fish. Soon there would be too few fish to feed local populations and no excess catches to sell for profit. But Lake Victoria is a very big lake, and the Nile perch is a very big fish (more than two meters long). A big fish in a big lake seemed like an ideal combination to attract commercial fishermen with big, elaborate nets from the outside world – right? Wrong. Native fishermen had been harvesting native fishes called cichlids, which eat mostly detritus and aquatic plants. The Nile perch eats other fish – including cichlids. Having had no prior evolutionary experience with the new predator, the 200 coexisting species of cichlids that were native to Lake Victoria had no defenses against it. And so the Nile perch ate its way through the cichlid populations and destroyed the lake’s biodiversity. Dozens of cichlid species found nowhere else are extinct. Without the cichlids to clean up the lake bottom, levels of dissolved oxygen plummeted and contributed to frequent fish kills. By 1990, fishermen were catching mostly Nile perch. Now there are signs that the Nile perch population is about to crash. By destroying its Kalafsky 40 food source, the Nile perch has undercut its own population growth. It has ceased to be a potentially large, exploitable food source for the people who live around the lake. As if that weren’t enough, the Nile perch is an oily fish. Unlike cichlids, which can be sun dried, the Nile perch must be preserved by smoking – and smoking requires firewood. Local people started cutting down more trees in the local forests, and trees are not rapidly renewable resources. To add insult to injury, the people living near Lake Victoria never liked to eat Nile perch anyway. They prefer the flavor and texture of cichlids. What is the lesson? A little knowledge and some simple experiments in a contained setting could have prevented the whole mess at Lake Victoria. THE RABBITS THAT ATE AUSTRALIA. During the 1800’s, British settlers in Australia just couldn’t bond with koalas and kangaroos, so they started to import familiar animals from their homeland. In 1859, in what would be the start of a wholesale disaster, a landowner in northern Australia imported and released two dozen wild European rabbits. Good food and good sport hunting, was the idea. An ideal rabbit habitat with no natural predators – that was the reality. Six years later, the landowner had killed 20,000 rabbits and was besieged by 20,000 more. The rabbits displaced livestock, even kangaroos. Now Australia has 200 to 300 million hippity-hopping through the southern half of the country. They overgraze perennial grasses in good times and strip bark from shrubs and trees during droughts. You know where they’ve been; they transform grasslands and scrublands into erode d deserts. They have been shot and poisoned. Their warrens have been plowed under, fumigated, and dynamited. Even when all-out assaults reduced their population size by 70 percent, the rapidly reproducing imports made a comeback in less than a year. Did the construction of a 2,000-mile-long fence protect western Australia? No. Rabbits made it to the other side before workers completed the fence. In 1951, government researchers introduced myxoma virus by way of mildly infected South American rabbits, its normal hosts. This virus causes myxomatosis. The disease has mild effects on the South American rabbits that coevolved with the virus but nearly always had lethal effects on the European rabbits. Biting insects, mainly mosquitoes and fleas, quickly transmit the virus from host to host. Having no coevolved defenses against the novel virus, the European rabbits died in droves. As you might expect, natural selection has since favored rapid growth of populations of the European rabbits that are resistant to the virus. In 1991, on an uninhabited island in Spencer Gulf, Australian researches released a population of rabbits that they had injected with a calicivirus. The rabbits died quickly and relatively painlessly from blood clots in their lungs, heart, and kidneys. In 1995, the test virus escaped from the island, possibly on insect vectors. It has been killing 80 to 95 percent of the adult rabbits in Australian regions. At this time, researchers are now questioning whether the calicivirus should be used on a widespread scale, whether it can Kalafsky 41 jump boundaries and infect animals other than rabbits (such as humans), and what the long-term consequences will be. THE PLANTS THAT ATE GEORGIA. A vine called kudzu (Pueraria lobata) was deliberately imported from Japan to the United States, where it faces no serious threats from herbivores, pathogens, or competitor plants. In temperate parts of Asia, kudzu is a well-behaved legume with a well-developed root system. It seemed like a good idea to import it for erosion control on hills and near highways in the southeastern United States. However, with nothing to stop it, kudzu’s shoots can grow one-third of a meter per day. Vines now blanket stream banks, trees, telephone poles, houses, hills, and almost everything else in their path. Attempts to dig them up or burn them are futile. Grazing goats and herbicides help, but goats are indiscriminate eaters and herbicides contaminate water supplies. If the global temperature continues to rise, kudzu could reach the Great Lakes by the year 2040. On the bright side, a Japanese firm is constructing a kudzu farm and processing plant in Alabama. Asians use a starch extract from kudzu in beverages, candy, and herbal medicines. The idea is to export the starch to Asia, where the demand currently exceeds supply. Also, kudzu might eventually help reduce the extent of logging operations. At the Georgia Institute of Technology, researches have reported that kudzu may be used as an alternative source of paper.