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Name: SN: Biology 413 (Zoogeography) Final Exam Winter Term 2 - 2015 Directions: 1. Write your name and student number on each page of the exam. 2. All answers should be written in the space provided. If you need extra space, you can use extra paper, but clearly label your answers. 3. The exam is designed to be completed in 2.5 hours. 4. The exam questions are organized into two parts and consist of seven pages. 5. PART I consists of 10 short-answer questions. 6. PART II consists of two long-answer (essay-form) questions and one multi-part question. Points PART I: 30 points PART II: 45 points TOTAL: 75 points PART I Answer the following 10 questions using 2-3 sentences or less. These questions should take you less than 5 minutes each. Please use the space provided. Each question in this section is worth 3 points. Question 1: Define the target effect and the rescue effect, in the context of the equilibrium theory of island biogeography. The target effect is the tendency for immigration rates to be higher on larger islands because larger islands have more shoreline, can be seen from farther away (i.e., create a larger target for colonization). The rescue effect is the tendency for extinction rates to be relatively low when islands are very close to the mainland because island populations are augmented by immigration. Question 2: What are the three dominant cycles that together make up the Milankovitch cycles? In general, what do these cycles impact to influence the Earth’s climate system and the advance and retreat of glaciers? Eccentricity (or ellipticity of orbit); Axial tilt (obliquity); Precession (pole wandering). The cycles generally impact the seasonality and location of solar energy around the Earth, thereby influencing Earth’s climate cycles and the advance and retreat of glaciers 1 Name: SN: Question 3: In addition to the Milankovitch cycles, what three related feedback mechanisms were associated with the growth/retreat of ice sheets and also contributed to rates of cooling and warming? With the growth and retreat of ice sheets, respectively, there was a 1) decrease and increase in plant and animal life over large areas, 2) decline and increase in production of greenhouse gases and 3) and increase and decrease in the amount of white surface on the earth, owing to an increase and decrease of the albedo effect. Question 4: Haffer’s (1969) hypothesis represents the “old view” that “islands” of Amazonian lowland rain forest persisted during the last glacial maxima and can be used to explain current patterns of disjunct taxa and speciation. The “new view” is that the Amazon did not exhibit isolated forest refugia during the Pleistocene. Provide one specific line of evidence that supports the new view. (Hint: we discussed two lines of evidence, one from botanical data and one from animal taxa). From botanical data: fossil pollen data from lake cores shows continuous forest cover and invasion by cold-adapted species during the last glacial maximum From animal taxa: range limits of trumpeter species in South America indicate separation by large river systems, not glacial refugia. The timing of diversification events shows that speciation of trumpeters occurred before the last glacial maxima. Question 5: Endemic species are sometimes described as “relicts”. What is the difference between taxonomic and biogeographic relicts? Taxonomic relicts are remnants of, at one time, a much more diverse taxon. Biogeographic relicts are taxa that at one time had a much wider geographic distribution. The difference emphasizes taxa that were once historically diverse (irrespective of distribution) versus taxa that were historically widespread (irrespective of diversity). Question 6: In the development of continental drift theory, many lines of evidence were used to better understand the mechanism by which continents shifted. We reviewed seven lines of evidence in lecture – one of these includes patterns of disjunct distributions in extant taxa. List three additional lines of evidence below. Can be any three of the following: 1) stratigraphic evidence (alignment of Precambrian shield and flood basalt deposits with Pangaea/Gondwana); 2) late Paleozoic glacial deposits are found on all continents of Southern Hemisphere/tongue fern fossil distribution and animal fossils are shared among now separate continents; 3) sea floor mapping; mid-Atlantic Ridge and other long chains of mid-ocean ridges suggests areas of plate shift; 4) paleomagnetism, magnetic stripes and magnetic reversals; certain metals are sensitive to polarity of earth’s magnetic field, alternating/symmatric patterns of polarity/direction on either side of ridges; 5) sea floor spreading; observations of magnetic striping and sea floor terrain suggested process of spreading; 6) concentration of earthquake activity associated with meeting/spreading of plates. Question 7: What are three potential causes of disjunction of populations or related species groups? (Hint: Continental drift represents a more specific case of one of these general potential causes.) Vicariance; extinction; long-distance dispersal of species (it’s okay to list these) ___________________ 2 Name: SN: Question 8: In the context of conserving biodiversity and evolutionary history, describe the basic motivation for conservation based on evolutionarily significant units (ESUs) and evolutionarily distinct and globally endangered (EDGE) species. ESUs are intraspecific groups that represent distinct characteristics and tendencies, occupy unique habitats or display unusual phenotypes. Phenotypic distinctiveness is often correlated with genetic distinctiveness. When considering conservation priorities, ESUs should be examined to capture an array of intraspecific diversity, and thus evolutionary potential for adaptation. EDGE species are recognized for the amount of unique evolutionary history that they represent, as well as their level of endangerment (e.g., based on the IUCN). For EDGE species, we ask how much evolutionary history would be lost if that species were to go extinct, as well as the likelihood of extinction given current and future threats. Question 9: What are the two metrics used to assess a mass extinction, and what to they measure? Extinction is measured by rate and magnitude. Rate measures the number of extinctions over a period of time in which the extinctions occurred (or the fraction of species that have gone extinct per unit time). Magnitude is the percentage of species that have gone extinct. ___________________________________________________________________________________ ___________________________________________________________________________________ ___________________________________________________________________________________ ___________________________________________________________________________________ ___________________________________________________________________________________ ___________________________________________________________________________________ Question 10: Provide a brief definition for the three “shortfalls” (Wallacean, Linnaean and Hutchinsonian) with respect to predicting changes in species distributions. The Wallacean shortfall refers to the deficit of lack of knowledge of explicit spatial distributions of species (i.e., where do species occur, what is the extent of distribution and patterns of density across a species range?). The Linnaean shortfall refers to shortcomings in the discovery and description of new species, or the deficit in knowing what species exist, as well as in classifying them as distinct from other species. The Hutchinsonian shortfall refers to our lack of knowledge of attributes of species niches and their interactions with the environment and other species, particularly in what species require for sustainable/persistent populations. 3 Name: SN: PART II Answer the following 3 questions using the space provided, with complete sentences as necessary. Each question in this section is worth 15 points. Question 11: We have discussed Janzen’s hypothesis with respect to several different themes in biogeography. Describe the predictions derived from Janzen’s hypothesis for tropical and temperate mountains and/or montane species with respect to the following: (1) physiological barriers to dispersal; (2) breadth of elevational ranges; (3) beta diversity (or change in species composition) along elevational gradients; (4) potential for allopatric speciation and (5) sensitivity of species to climate warming. [3 points each – 15 points total] (1) Janzen’s hypothesis predicts that mountains are “higher” in the tropics. Specifically, species occurring in the tropics experience and are specialized to a narrower range of thermal conditions annually compared to temperate species. Therefore a mountain of a given height in the tropics would pose a greater barrier to dispersal for a species compared to a mountain in the temperate zone, because a given tropical species would likely need to disperse through regions outside of its physiological tolerance to colonize new areas beyond mountain passes, more so than species in temperate regions, which would be less likely to encounter thermal regimes outside of its physiological range of tolerance. (2) A direct consequence of the narrower physiological tolerances of species in the tropics should be that tropical species also show narrower elevational ranges compared to temperate species. (3) If species, on average, have narrower elevational ranges in the tropics, then we would expect beta diversity (or spatial species turnover, or change in species composition at the community level) to be higher in the tropics compared to the temperate zone, where species on average have broader elevational ranges. (4) With mountains being more significant dispersal barriers and with species having narrower elevational ranges in the tropics, we would expect the potential for allopatric speciation to also be higher in the tropics, compared to the temperate zone, simply because we would expect more opportunity for isolation, either through vicariance, or through infrequent successful dispersal across barriers. (5) Following Janzen’s hypothesis, if species have narrow physiological tolerances in the tropics, then any incremental change in temperature may bring species closer to their physiological limit, compared to species in temperate areas with broader physiological tolerances. Therefore, we would expect tropical species to be more sensitive to a similar magnitude of climate warming, compared to temperate species. This hypothesis has been supported in various species of ectotherms, including insects and lizards. 4 Name: SN: PART II (continued) Question 12: Relying on what you know about the Theory of Island Biogeography and the information provided in the questions, reconstruct the box exhibiting MacArthur and Wilson’s Immigration/Extinction dynamics. [15 points total, including parts a - d] (a) The empty box below is labeled with arrows that represent increasing isolation on the x-axis and increasing area on the y-axis. At the top of the box (upper portion of the box opposite the x-axis) indicate one of two rates associated with increasing isolation (immigration or extinction) with an arrow showing the rate as increasing. Draw another arrow on the right-hand side of the graph (opposite the y-axis) indicating the other rate (immigration or extinction) showing the rate as increasing (2 points). (b) The following three terms can be used to describe the composition of the communities found in each corner of the box. Match one term to each corner by writing the term in the box, based on the axes provided and the axes you created. The term for the lower right-hand corner is provided (3 points). High endemism Mainland like Species poor/Highly transient (c) The following terms describe the processes that largely shape the composition of communities in three corners of the box you filled in above. Indicate which process is associated with each corner, based on the axes that are provided and the axes you have created in part (a) above (3 points). Evolution Stochastic events/neutrality Ecological interactions 5 Name: SN: PART II (continued) (d) Now, put your ideas together to explain the box you reconstructed above. For each of the three corners that you filled in, explain why we see predictable patterns in composition within each community (part b), which are generally shaped by each process (part c). Also explain why we see depauperate islands in the lower right-hand corner. Be sure to relate the terms in each corner to both the area/isolation axes provided and the immigration/extinction rate axes you created in part (a). Use complete sentences (7 points). Top left corner: With islands that are large in size and close to the mainland (low isolation), we expect the composition of communities to be more representative of mainland communities, due to higher rates of immigration/exchange of individuals and low rates of extinction. Thus, the processes that largely govern the composition of these communities are ecological in nature (lots of species are present to interact, plus there is less opportunity for these species to be isolated for any long period of time; selection is less effective in the face of high gene flow). Top right corner: These islands will experience lower rates of colonization due to high isolation, but also lower rates of extinction due to larger size. As such, the species that are able to reach the island are more likely to persist there over time. Without constant gene flow from the mainland, these communities are more subject to evolutionary pressures like selection, and such islands, given enough time should host high numbers of (in-situ) endemic species, showing communities that are more distinct from the mainland. Bottom left corner: Because these islands are small, they will experience higher extinction rates, but due to their close proximity to the mainland, they will be more readily colonized by individuals dispersing from the mainland. The resulting community should be one that is relatively species poor, because the island is small and cannot support many species overall, but also one that is also highly transient and subject to stochastic events, due to continual/repeated colonization and higher extinction. (Ecological interactions are not as important in determining species composition of these islands, because the transient nature of the community with high colonization/extinction rates leads to unpredictable dynamics of populations for most species). Bottom right corner: these islands are both small in area and far from the mainland. We expect few species to be able to reach these islands in the first place (low colonization), and because the islands are small, they experience higher extinction rates. This results in species poor/depauperate faunas overall. 6 Name: SN: PART II (continued) Question 13: The latitudinal gradient in species diversity is sometimes considered the only true law in biogeography. Explain what this gradient is (3 points) and provide a discussion of four major hypotheses to account for it (including their weaknesses or exceptions to the general patterns; 3 points per hypothesis). We discussed more than four hypotheses in class, but choose four among those presented. You may use illustrations to help make your points. [15 points total] The latitudinal gradient in species diversity is the well-established pattern that areas/habitats at lower latitudes (tropical regions, towards the equator) tends to be higher in species richness than areas/habitats at higher latitudes (temperate regions, towards the poles). The hypotheses can be generally categorized into null hypotheses, deterministic/non-equilibrial and deterministic/equilibrial hypotheses. Looking for specific hypotheses within these broad categories. Null hypotheses: (1) Mid-domain effect. Given any bounded gradient, such as a latitudinal gradient, we would expect the greatest range of species overlap at a mid-point between the extremes of the gradient, or in the case of latitude, mid-way between the poles. Thus the pattern is an artifact with no underlying biological significance. Deterministic, non-equilibrial hypotheses: (2) Communities have not reached a steady state; communities, and species diversity is still in the process of increasing (or decreasing) following historical disturbance or dynamics. Deterministic, equilibrial hypotheses (assumes a steady state has been reached): (3) Higher productivity (i.e., solar radiation, potential evapotranspiration) in the tropics leads to higher resource availability, such that species can more finely partition a resource gradient (increased species packing). Exceptions to this include shallow eutrophic lakes and salt marshes, which are extremely productive, but have low species diversity. (4) Harsh environments, characteristic of the temperate zone (e.g., icy and cold), have higher extinction rates, lower colonization potential, and less opportunity for resource specialization than more benign environments. (5) Temporal stability is higher in the tropics. Variable climates prevent resource specialization, and hence are able to support fewer species. Temporal variability tends to favour generalists at the expense of specialists. Exceptions include the deep ocean, which is stable, but has low diversity (but also lower productivity). (6) Tropical habitats are more structurally diverse, with higher habitat heterogeneity. Diverse physical environments promote isolation, resource specialization, speciation, and co-existence. Exceptions include marine plankton communities, which are highly diverse despite low heterogeneity. (7) Interspecific Interactions (and feedback). More species in tropical communities create a positive feedback of diversification through increased competition, predation, and parasitism. Not certain if this is a cause for tropical diversity or a consequence of it (chicken – egg). (8) Area. Tropics have greater area than either polar region due to curvature of the earth. Species-area relationship is well established. However, tropical latitudes have much less terrestrial area. (9) Age. Latitudinal gradients for clades originating in warm climates are steeper, with a strong tropical affinity. For a variety of plants and animals of both marine and terrestrial realms, most clades radiated in tropical climates. Extant tropical diversity peak is created from lineages that adapted to a planet with tropical climate. Higher diversities have arisen among tropical clades because the earth has been predominantly tropical throughout most of its history. 7