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91605 916050 3SUPERVISOR S Level 3 Biology, 2016 91605 Demonstrate understanding of evolutionary processes leading to speciation 2.00 p.m. Thursday 10 November 2016 Credits: Four Achievement Achievement with Merit Achievement with Excellence Demonstrate understanding of evolutionary processes leading to speciation. Demonstrate in-depth understanding of evolutionary processes leading to speciation. Demonstrate comprehensive understanding of evolutionary processes leading to speciation. Check that the National Student Number (NSN) on your admission slip is the same as the number at the top of this page. You should attempt ALL the questions in this booklet. If you need more room for any answer, use the extra space provided at the back of this booklet and clearly number the question. Check that this booklet has pages 2 12 in the correct order and that none of these pages is blank. YOU MUST HAND THIS BOOKLET TO THE SUPERVISOR AT THE END OF THE EXAMINATION. TOTAL New Zealand Qualifications Authority, 2016. All rights reserved. No part of this publication may be reproduced by any means without the prior permission of the New Zealand Qualifications Authority.

2 QUESTION ONE: MEXICAN SPADEFOOT TOAD The Mexican spadefoot toad (Spea multiplicata) is found in southwestern United States and Mexico. In ponds with low abundance of food resources and high density levels of tadpoles, two populations predominate. One population (called the omnivore morph) has a round body with a long intestine, small jaw muscles, smooth mouth parts, and has a generalist omnivorous diet of algae and small crustaceans found on the bottom of the pond. The other population (called the carnivore morph) has a narrow body with a short intestine, enlarged jaw muscles, teeth-like mouthparts, and has a specialist carnivorous diet of fairy shrimps found in the water column. On the other hand, in ponds of high abundance of food resources and low density levels of tadpoles, only one population, of intermediate phenotype, is found. Compare and contrast the impact of disruptive and stabilising selection on genetic diversity AND discuss how speciation could occur in the Mexican spadefoot toad. In your answer you should: describe genetic variation Figure 1: Mexican spadefoot toad tadpoles from a high density, low food resource pond. Top: the omnivore morph. Bottom: the carnivore morph. http://labs.bio.unc.edu/pfennig/labsite/photos.html describe the terms disruptive and stabilising selection, and describe which population(s) of Mexican spadefoot toad tadpole is associated with each type of selection explain the selection pressures that promote disruptive selection, AND the selection pressures that promote stabilising selection in the Mexican spadefoot toad tadpole. Well labelled diagrams can be used to support your answer.

3 There is more space for your answer to this question on the following page.

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5 QUESTION TWO: THREE-SPINED STICKLEBACK The three-spined stickleback (Gasterosteus aculeatus) is a small (30 90 mm) fish found in the Northern Hemisphere. Some populations live in coastal marine habitats, while other populations live in freshwater. Three-spined sticklebacks lack the scales typical of most fishes; instead they possess (protective) bony plates and spines. Three-spined stickleback populations living in a marine habitat have high numbers of bony plates and long spines, whereas freshwater populations typically have low numbers of bony plates and short spines. Genetic evidence suggests that a mutation in the Ectodysplasin (EDA) gene causes variation in plate number, and a mutation in the PITX1 gene causes variation in spine length. The main predators of three-spined stickleback in marine habitats are larger fish. In freshwater habitats, grasping insects (such as dragonfly larvae) are the main predators, especially of juvenile three-spined stickleback. Marine habitats typically have low amounts of shelter suitable for the three-spined stickleback, whereas freshwater habitats have high amounts of shelter. The growth rate and acceleration / burst speed of three-spined sticklebacks is highest when the bony plate number is lowest. Discuss how EDA and PITX1 gene mutations AND natural selection have affected evolution in three-spined stickleback. Figure 2. Top: Typical three-spined stickleback from a marine population. Bottom: Typical three-spined stickleback from a freshwater population. Fish have been stained with alizarin red to highlight bony plates and spines. http://unews.utah.edu/wp-content/uploads/ sticklebackfigure1.jpg Figure 3. Typical three-spined stickleback predators in ocean and freshwater habitats. http://learn.genetics.utah.edu/content/selection/stickleback/ In your answer you should: describe the terms mutation AND natural selection explain how selection pressures in marine AND freshwater habitats act differently on bony plate number and spine length discuss the roles of mutation AND natural selection on three-spined stickleback evolution. There is more space for your answer to this question on the following page.

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8 QUESTION THREE: KAKARIKI Kakariki are the most common species of parakeet in the genus Cyanoramphus and are distributed throughout the South Pacific (Figure 5). Aotearoa has the largest number of species. Kakariki live in a wide range of habitats, including subantarctic tussock (Antipodes Island kakariki and Reischek s kakariki), beech forests in mainland Aoteoroa (yellow-crowned kakariki and orange-fronted kakariki), and tropical rainforests (New Caledonian red-crowned kakariki). Figure 4. Forbes kakariki, Chatham Island. www.nzbirdsonline.org.nz/species/forbes-parakeet New Caledonia New Caledonian red-crowned kakariki Society Islands Society Island kakariki (extinct) Black-fronted kakariki (extinct) Norfolk Island Norfolk Island kakariki Kermadec Islands Kermadec kakariki Lord Howe Island Lord Howe Island kakariki (extinct) Stewart Island Red-crowned kakariki Auckland Islands Red-crowned kakariki Macquarie Island Macquarie Island kakariki (extinct) North Island Yellow-crowned kakariki Red-crowned kakariki South Island Yellow-crowned kakariki Red-crowned kakariki Orange-fronted kakariki Chatham Islands Forbes kakariki Chatham Island red-crowned kakariki Antipodes Islands Antipodes Island kakariki Reischek s kakariki Campbell Island Campbell Island kakariki (fossil evidence) Figure 5: Kakariki distribution in the South Pacific.

The evolutionary relationships of kakariki species have been determined using mitochondrial DNA sequence analysis. The phylogenetic tree based on this analysis is shown in Figure 6. The climate during this period is shown in Figure 7, and the reconstructed vegetation cover at the height of the last glacial period is shown in Figure 8. 9 Chatham Island red-crowned kakariki Red-crowned kakariki Reischek s kakariki Orange-fronted kakariki Yellow-crowned kakariki Norfolk Island kakariki Antipodes Island kakariki Forbes kakariki New Caledonian red-crowned kakariki 500 000 400 000 300 000 200 000 100 000 0 Years before present Figure 6. Phylogenetic tree for Cyanoramphus. The time scale for evolutionary divergence is indicated above. Adapted from Boon, W. M. et al. (2001). Molecular systematics and conservation of the kakariki (Cyanoramphus spp.), Science for Conservation, 176 (Department of Conservation, Wellington). Years before present day 0 14 500 75 000 125 000 180 000 220 000 280 000 320 000 370 000 450 000 500 000 Cool period Warm period Figure 7. Glacial periods in Aotearoa. Adapted from www.teara.govt.nz/en/ diagram/10741/glacial-periods-in-new-zealand Figure 8. Aotearoa vegetation cover 19000 29000 years b. p. as reconstructed from pollen, macrofossil, beetle and geographic evidence. Adapted from: Newnham, R, et al. (2010). The vegetation cover of New Zealand during the last glacial maximum, terra australis, 32, p. 59 (ANU E Press, Canberra). http://press.anu. edu.au/wp-content/uploads/2011/02/ch0417.pdf

10 Discuss the pattern of evolution in kakariki, and the factors that have affected kakariki evolution. In your answer you should: describe the evolutionary pattern AND type of speciation indicated by the resource material explain the origin and distribution of kakariki in Aotearoa with reference to the phylogenetic tree using the information provided, discuss how biological and geographical factors have contributed to kakariki speciation.

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12 QUESTION NUMBER Extra paper if required. Write the question number(s) if applicable. 91605