Adaptive Radiation in Rain Forest Birds

Activity #2 Rain Forest Unit 3 Activity #2 Adaptive Radiation in Rain Forest Birds

In Advance Student Reading Assign the Student Page Adaptive Radiation in Hawaiian Honeycreepers (pp. 22-26) as homework. Class Period One Adaptive Radiation Discussion Materials & Setup For each student Student Page Adaptive Radiation in Hawaiian Honeycreepers (pp. 22-26) Student Page Adaptive Radiation in Hawaiian Honeycreepers: Questions on the Reading (pp. 27-28)

Instructions 1) Have students complete the Student Page Adaptive Radiation in Hawaiian Honeycreepers: Ques- tions on the Reading in class.

2) Spend the remainder of the class discussing adaptive radiation and the homework reading, begin- ning with student responses to the questions. Journal Ideas Many native birds are not found in low-elevation areas on Maui. Do you think that native birds once occupied these lowland areas? How would you go about finding out? Do you think anything should be done to protect the remaining Hawaiian honeycreepers? Why or why not? Keeping in mind the different human-caused pressures that have led to declines and extinctions among Hawaiian honeycreepers, what do you think can be done to protect the species that remain? Imagine being a traditional Hawaiian bird catcher, collecting thousands of feathers over your lifetime. What do you think it would have been like to work mostly alone in the forests of the gods? Assessment Tools Student Page Adaptive Radiation in Hawaiian Honeycreepers: Questions on the Reading (teacher version, pp. 20-21) Participation in class discussion Journal entries

Rain Forest Birds: A Study in Adaptation Höÿike o Haleakalä 19 Activity #2 Rain Forest Unit 3

Teacher Version Adaptive Radiation in Hawaiian Honeycreepers: Ques- tions on the Reading

1) Define adaptive radiation, and explain its relationship to endemic species. Give one example of adaptive radiation in Hawaiian species other than honeycreepers.

Adaptive radiation is the evolution of many species from a single ancestor.

Well-reasoned responses about the relationship between adaptive radiation and endemism are acceptable. Possible relationships include: Adaptive radiation results in many endemic species (species that evolved here and are found nowhere else in the world) that are closely related to each other because of their common ancestor, and Because species are evolving in response to local conditions, adaptive radiation may result in species that are narrowly endemic, or restricted to a small range or a single island.

Examples include Hawaiian Drosophila flies and Hawaiian lobeliads. There are many others that students may have learned about outside this unit, including the silversword alliance which includes the ÿähinahina or Haleakalä silversword.

2) Why are fossil records valuable sources of information to scientists studying the evolution of native bird species?

Well-reasoned responses are acceptable. Fossil records enable scientists to identify previously unknown species and establish their relationships with existing species. Among Hawaiian honeycreepers, for example, 18 species are known only from fossil records. Thats 35 percent of all known honeycreeper species. Our understanding of the scope of adaptive radiation among honeycreepers would be much narrower if not for the fossil record.

20 Rain Forest Birds: A Study in Adaptation Höÿike o Haleakalä Activity #2 Rain Forest Unit 3

3) Is the shift in the size of ÿiÿiwi bills over the last 100 years an example of adaptive radiation in action? Explain your answer.

Well-reasoned responses are acceptable. Two possibilities: The shift in ÿiÿiwi bill size is probably not adaptive radiation in action because it is likely to be taking place across the entire species, so the changes that are taking place are probably not going to result in the creation of a new species. The extinction of the ÿöÿö probably affected the ÿiÿiwi across much of its range, as did the decline of the preferred food source, Hawaiian lobeliads. If there are islands or large stretches of habitat in which Hawaiian lobeliad populations are protected or restored and other places where they are not, then we may be seeing adaptive radiation in action, as the habitat for some populations of ÿiÿiwi would favor their existing bill size, and the habitat for others would favor a shorter bill, possibly leading to species differentiation over a long period of time.

4) Using what you have learned about evolution and adaptation, explain why extinctions of rain forest bird species have happenedand continue to occurso rapidly in the face of human-caused changes to native Hawaiian rain forests.

Well-reasoned responses are acceptable. Two possibilities: Human-caused changes such as habitat destruction, pressures by introduced species, and introduced diseases are altering conditions for native birds so quickly that evolution cannot keep up. The honeycreepers took millions of years to evolve, but humans have caused dramatic changes in their environment within hundreds of years. Human activity has contributed to many different pressures on native birds including habitat destruction, predation and competition by introduced species, and disease. A species that might be able to adapt and survive in the face of a single human-caused pressure may not fare so well when there are multiple pressures working against its survival.

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Adaptive Radiation in Hawaiian 123456

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The Hawaiian Islands are the most isolated away. These new islands may have offered 123456

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123456 archipelago on the planet. Here, we are more than different habitat for the birds, which over time 123456 123456

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123456 2,000 miles away from the nearest continent and could have further radiated into new species. 123456 123456

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123456 some 1,000 miles away from the closest Pacific In the 1970s, paleontologists began in earnest 123456

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123456 atolls. Because of this isolation, over millions of to collect partially fossilized bird remains from 123456

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123456 years, relatively few plant and animal species sand dunes, lava tubes, caves, and sinkholes in 123456 123456

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123456 arrived on the islands. The three Ws (wind, the Hawaiian Islands. This fossil record revealed 123456 123456

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123456 waves, and wings) are used to describe the means many species of honeycreepers that were already 123456

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123456 by which species arrived here. From a small 123456 123456

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123456 collection of founding species descended 123456 123456

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123456 all of the animals and plants that are native 123456

Hawaiian Honeycreepers 123456

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123456 to Hawaiÿi. The Hawaiian rain forest is a 123456

123456 at a Glance 123456 123456 hotbed of endemism. Of its native inhabit- 123456 123456

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123456 ants, a large proportion are unique to these 123456 123456

123456 Thought to have evolved from a single spe- 123456 123456 islands. 123456

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cies of finch. 123456 In contrast to indigenous species, 123456 123456

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123456 Hawaiian endemic species evolved into a 123456 123456

123456 Fifty-two species are thought to be part of the 123456 123456 new species after arriving here from some- 123456

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Hawaiian honeycreeper subfamily. 123456 where else. Indigenous species have re- 123456

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Eighteen species, known only 123456 mained relatively unchanged since their 123456 123456

123456 from fossil record, were 123456 arrival and are not unique to Hawaiÿi. 123456 123456

123456 probably extinct by 1778. 35 percent 123456 123456

Among the endemic forest birds, most 123456

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Twelve species have gone 123456 species belong to a group known as the 123456 123456

123456 extinct since 1778. 23 percent 123456 Hawaiian honeycreepers. All of the honey- 123456 123456

123456 Fourteen surviving species 123456 123456 creeper species are thought to have evolved 123456

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are classified as endangered, 123456 from a single finch species that arrived on 123456

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some may already be extinct. 27 percent 123456 the islands more than 15 million years ago. 123456 123456

123456 Eight surviving species are 123456 This process of adaptive radiation has, 123456 123456

123456 not classified as endangered. 15 percent 123456 123456 over time, resulted in honeycreepers that are 123456

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123456 very different from each other and adapted 123456 123456

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123456 to many different environments. 123456 123456

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The oldest of todays main islands, Kauaÿi, is extinct by the time Western scientists had started 123456

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123456 only about five million years old. So much of the identifying and describing Hawaiian birds. Most 123456

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123456 early evolution of the honeycreepers probably of these extinctions were most likely a result of 123456 123456

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123456 happened on other, older islands in the chain. the impacts of the Hawaiian people: competition 123456 123456

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Through a process called interisland dispersal, or predation by Polynesian introductions, conver- 123456

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123456 the ancestors of the honeycreeper species we sion of lowland habitats into agricultural areas, 123456 123456

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123456 know today colonized the new islands, even as and killing of birds for food and other human 123456 123456

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123456 their old homelands were being slowly eroded uses. 123456

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123456 In addition to uncovering evidence of bird Their beak structure and even their leg and facial 123456 123456 123456

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123456 extinctions, fossil research has helped to shed muscles reflect their diet and how they forage for 123456

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123456 new light on the true extent of adaptive radiation food. 123456 123456

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123456 2 of 5 of 2 in Hawaiian honeycreepers. From the single Some nectar-eating honeycreepers have beaks 123456 123456

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123456 ancestral finch species, at least 52 honeycreeper that are shaped to fit specific flowers. This is 123456

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123456 species are known to have evolved. What caused evidence of coevolution, in which honeycreepers 123456

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123456 this remarkable formation of new species? and flowering plants evolved together, the birds 123456 123456

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123456 feeding on the nectar and in turn spreading pollen 123456 123456

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from flower to flower. Birds and flowers evolved 123456

Adaptive Radiation 123456

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123456 specialized relationships, of which the ÿiÿiwi and 123456 Adaptive radiation is often thought of as being 123456 123456

123456 Hawaiian lobeliad plants are an excellent ex- 123456 123456 driven by the need or opportunity for plants or 123456

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ample. The long, curved bill of the ÿiÿiwi is 123456 animals to live in habitats other than the ones to 123456 123456

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perfectly suited to gathering nectar from the long, 123456 which they are best adapted. It is easy to see how 123456 123456

123456 tubular, curved flowers of many lobeliad species. 123456 123456 such needs and opportunities could arise for birds 123456

123456 Many factors are thought to have influenced 123456 123456 arriving on the Hawaiian Islands from other parts 123456

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adaptive radiation in Hawaiian honeycreepers, 123456 of the world. The Hawaiian honeycreepers 123456 123456

123456 including the following: 123456 evolved into species with different primary food 123456 123456

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123456 sources: nectar, hard seeds, soft fruit, and insects 123456

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123456 Isolation and Dispersability 123456 and larvae. Some honeycreeper species have diets 123456 123456

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123456 made up of combinations of these food sources. Populations of a bird species that remain 123456 123456

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isolated from each other can, over time, evolve 123456

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into separate species. In general, birds can easily 123456

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123456 disperse from one placeor island to another 123456 123456

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123456 by flying. But the pattern of species and island- 123456 123456

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specific subspecies of honeycreepers suggests 123456

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123456 that some species were more likely to do so than 123456 123456

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123456 were others. 123456 123456

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Some scientists believe that insect-eating 123456

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honeycreepers have a more reliable source of 123456

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123456 food than do species that feed on fruit or nectar. 123456 123456

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123456 They hypothesize that frugivorous (fruit- 123456 123456

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eating) and nectivorous (nectar-feeding) birds 123456

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123456 are more likely to fly between islands looking for 123456 123456

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123456 food in lean years. Because the insectivores 123456 123456

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(insect-feeders) had less reason to move around, 123456

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they remained more isolated and developed into 123456

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123456 more species and subspecies. 123456 123456

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123456 Other scientists believe that there is not such a 123456 123456

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clear link between eating habits and interisland 123456

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123456 dispersal. Instead, they distinguish between 123456 123456

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123456 common generalists such as the ÿapapane and 123456 123456

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ÿamakihi that are widespread across the islands 123456

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on the one hand, and specialized seed eaters and 123456

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123456 Variations in bill morphology of selected nectivores on the other hand. Generalists are able 123456 123456

123456 Hawaiian honeycreepers 123456 to feed on a variety of food types. The more 123456 123456

123456 (Illustration: H. Douglas Pratt, Jr.) 123456 123456

specialized eaters, some scientists maintain, 123456

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would have been more likely to stay where their 123456

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123456 food source was and further evolve in that place. 123456 123456

Other Rain Forest Examples of 123456

123456 The generalists would have been more likely to 123456 123456

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Adaptive Radiation 5 of 3 123456

fly from island to island. 123456

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123456 The Hawaiian Drosophila flies: Over 500 123456 123456 Adaptation to Food Resources 123456 123456

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species of flies in the Drosophilidae 123456

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Birds can rapidly adapt to new types of food or 123456

123456 family have been identified in Hawaiÿi, all 123456 123456 changes in the abundance of food sources. Over 123456 123456 of which evolved from a single common 123456 123456 time, these adaptations (physical or behavioral 123456 123456 ancestor species. Nearly one-quarter of the 123456 123456

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changes) can result in the emergence of new 123456

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known species in this family are found in 123456

123456 species. In 1984 and 1985, studies of endangered 123456 123456 Hawaiÿi, including many that are nar- 123456 123456 Laysan finches that had been introduced in 1967 123456 123456 rowly endemic. That means that they 123456 123456

123456 to Pearl and Hermes Reef, showed significant 123456

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occur in only a very small area. 123456

123456 differences had already developed between the 123456

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123456 Pearl and Hermes finches and their ancestors on 123456 123456 Hawaiian lobeliads: Six of seven genera of 123456 123456 Laysan. The differences appeared to be a re- 123456 123456 this group of plants are endemic to 123456 123456

123456 sponse to differences in food availability. If those 123456

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Hawaiÿi. The 98 species in the genera 123456

123456 differences turn out to be genetically linked, then 123456 123456 Cyanea, Clermontia, Delissea, and 123456 123456 evolutionary change had occurred among these 123456 123456 Rollandia are considered by many scien- 123456 123456

123456 birds in fewer than 20 generations. 123456

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tists to be the largest group of Hawaiian 123456

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123456 plants to have evolved from a single 123456 123456

123456 Intraspecies Food Competition 123456 immigrant species. 123456 123456

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Competition for food among members of the 123456

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123456 same species can lead to adaptive shifts or 123456 123456

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123456 changes. Less competitive members of the 123456 123456

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123456 Evolution is Not Over 123456 species may shift to feeding on a different size or 123456

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123456 type of food, and over time this division of food Evolutionary changes in Hawaiian honeycreep- 123456 123456

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123456 can lead to bills that are specialized to this avail- ers did not end at some time in the past. In fact, 123456 123456

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123456 there is evidence that rapid evolutionary changes 123456 able food source. 123456

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have occurred within the past 100 years, in 123456

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response to human-induced extinctions and 123456

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123456 habitat changes. Given the fact that the honey- 123456 123456

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123456 creepers evolved over millions of years, to mea- 123456 123456

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sure significant changes in one hundred years 123456

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123456 indicates rapid natural selection. 123456 123456

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123456 Here is an example of such a rapid shift: 123456 123456

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In the early 1990s, a group of biologists com- 123456

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pared body measurements of liveÿiÿiwi with 123456

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123456 museum specimens collected prior to 1900 123456 123456

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123456 (Smith, et al., 1995). Early studies of the ÿiÿiwi 123456 123456

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prior to 1900reported that the long, curved 123456

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123456 flowers of Hawaiian lobeliads were this birds 123456 123456

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123456 preferred food. Now, however, ÿiÿiwi feed mainly 123456 123456

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on the open flowers of ÿöhiÿa, which do not have 123456

123456 ÿIÿiwi (Photo: Eric Nishibayashi) 123456 123456

tubular, curved flowers. 123456

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123456 What might have caused this change in diet? changes open the possibility for rapid evolution- 123456 123456

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123456 Researchers point to two factors. The first is that ary shifts, such as the one that appears to be 123456 123456

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the lobeliads, once a prominent part of the under- happening with the ÿiÿiwi. 123456

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123456 4 of 5 of 4 story of Hawaiian rain forests, are now rare. 123456 123456

123456 th 123456 During the 20 century, habitat degradation and 123456 Ongoing Threats 123456 123456

123456 grazing by feral ungulates induced the extinction 123456 123456 Of the 22 known surviving species of Hawaiian 123456 123456

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of 25 percent of Hawaiian lobeliad species. Most 123456 honeycreepers, 14 are considered endangered, 123456 123456

123456 of the species that remain are rare or endangered. 123456 123456

and some of these may already be extinct. What 123456

123456 So the preferred food of the ÿiÿiwi is greatly 123456 123456

is contributing to this loss of species today? 123456

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reduced. 123456

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123456 Researchers suggest the second factor influenc- 123456 Habitat destruction: The Maui parrotbill is an 123456 123456

123456 ing this dietary shift was the extinction of the 123456 123456

illustrative example of the impact of habitat 123456

123456 ÿöÿö. The ÿöÿö was a Hawaiian honeyeater, 123456 123456 destruction. By the 1890s, it was already rare 123456 123456

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another native Hawaiian bird not related to the 123456 in the remaining koa-dominated rain forests. 123456 123456

123456 honeycreepers. This bird was behaviorally domi- 123456 123456

These koa forests were destroyed, largely by 123456

123456 nant over the ÿiÿiwi, keeping the ÿiÿiwi from 123456 123456

wildfires, browsing by introduced animals, 123456

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123456 feeding heavily in the favored trees of the ÿöÿö: 123456 and logging. Today, the parrotbill survives 123456 123456

123456 the ÿöhiÿa. By 1900, the ÿöÿö was extinct, and as 123456 only in higher-elevation ÿöhiÿa forests, where 123456 123456

123456 the lobeliads declined over the course of the 123456 123456

it is an extremely rare bird. 123456

123456 1900s, theÿiÿiwi shifted its foraging emphasis to 123456 123456

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ÿöhiÿa flowers. 123456 Avian malaria and other diseases: Like many 123456 123456

123456 Birds that feed most efficiently on ÿöhiÿa 123456 123456

Hawaiian honeycreepers, the rare ÿäkohekohe 123456

123456 flowers, such as the ÿapapane and the 123456 123456

is thought to be highly susceptible to avian 123456

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ÿäkohekohe, have short bills to exploit this food 123456 malaria, a disease introduced early this 123456 123456

123456 source. Researchers hypothesized that a shift in 123456 century with nonnative birds. The disease is 123456 123456

123456 diet away from the long, tubular, curved lobeliad 123456 123456

transmitted by a nonnative mosquito and is 123456

123456 flowers to the open ÿöhiÿa flowers would have 123456 123456 believed to prevent honeycreeper populations 123456 123456

123456 resulted in selective pressures which favored 123456 from surviving in lower-elevation forests. 123456 123456

123456 shorter bills over longer bills. 123456 123456

Mosquitos rarely occur above 1200 meters 123456

123456 The biologists tested this hypothesis by com- 123456 123456

(3936 feet) elevation. Many native birds 123456

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123456 paring bill measurements and other body mea- 123456 escape infection only while they remain in 123456 123456

123456 surements of live ÿiÿiwi with museum specimens 123456 upper-elevation forests. 123456 123456

123456 collected prior to the extinction of the ÿöÿö. 123456 123456

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123456 Evaluating the data they collected, they found 123456 123456 Predation by cats, mongooses, and rats: These 123456 123456

123456 that the upper mandible (beak portion) became 123456 animals eat native bird eggs and chicks, and 123456 123456

123456 shorter, while overall body size stayed the same. 123456 123456

sometimes even adult birds, especially those 123456

123456 The results of their analysis suggest the longer 123456 123456

tending their young. 123456

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123456 billed birds, presumably due to lower feeding 123456

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123456 effiencies on ÿöhiÿa, were lost over time as 123456 These pressures are happening at a rapid rate, 123456 123456

123456 selection favored shorter and possibly straighter 123456 123456

much more quickly than many bird species can 123456

123456 bills. 123456 123456 adapt. The honeycreepers that are faring the best 123456 123456

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As human-induced extinctions and declines of 123456 tend to be those that can adapt to a broad range of 123456 123456

123456 native species continue, the structure of Hawaiian 123456 123456

habitats. The most common honeycreepers on 123456

123456 natural communities is changing rapidly. These 123456 123456

Maui and in Hawaiÿi are the ÿapapane and the 123456

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123456 ÿamakihi. Both of these birds range from the rain 123456 123456

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123456 forests into upper-elevation shrublands and even 123456 123456

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into planted forests. As the ÿiÿiwi shows, however, 123456

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123456 rapid evolutionary shifts are possible, and that 123456 123456

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123456 5 of 5 of 5 possibilityalong with efforts at habitat protec- 123456 123456

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123456 tion, feral animal control, and research into avian 123456

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123456 diseasesmay spell survival for some of the 123456

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123456 remaining Hawaiian honeycreeper species. 123456 123456

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123456 Sources 123456 123456

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123456 Freed, L. A., Extinction and Endangerment of Juvik, James O., Biogeography, in Juvik, Sonia 123456 123456

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123456 Hawaiian Honeycreepers: A Comparative Ap- P., and James O. Juvik (eds), The Atlas of 123456 123456

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proach, in Landweber, L. F., and A. P. Dobson Hawaiÿi, 3rd ed., University of Hawaiÿi Press, 123456

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123456 (eds.), Genetics and the Extinction of Species, Honolulu, 1998, pp. 103-106. 123456 123456

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123456 Princeton University Press, Princeton, New 123456 123456

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123456 Jersey, 1999, pp. 137-162. 123456 Medeiros, Arthur C., and Lloyd L. Loope, Rare 123456

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Animals and Plants of Haleakala National Park, 123456

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123456 Freed, L. A., S. Conant, and R. C. Fleisher, Hawaiÿi Natural History Association, Hawaiÿi 123456 123456

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123456 Evolutionary Ecology and Radiation of Hawai- National Park, Hawaiÿi, 1994. 123456 123456

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ian Passerine Birds, in Kay, E. A. (ed.), A Natu- 123456

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123456 ral History of the Hawaiian Islands, Selected Smith, T. B., L. A. Freed, J. K. Lepson, and J. H. 123456 123456

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123456 Readings II, University of Hawaiÿi Press. Hono- Carothers, Evolutionary Consequences of 123456 123456

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123456 lulu, 1994, pp. 335-345. 123456 Extinctions in Populations of a Hawaiian Honey- 123456

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creeper, Conservation Biology, Vol. 9, No. 1, 123456

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123456 Givnish, T. J., K. J. Sytsma, J. F. Smith, and W. J. 1995, pp. 107-113. 123456 123456

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123456 Hahn, Molecular Evolution, Adaptive Radiation, 123456 123456

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and Geographic Speciation in Cyanea 123456

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(Campanulaceae, Lobelioideae), in Wagner, 123456

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123456 W. L., and V. A. Funk (eds.), Hawaiian Biogeog- 123456 123456

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123456 raphy: Evolution on a Hot Spot Archipelago, 123456 123456

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Smithsonian Institution, Washington, DC, 1995, 123456

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123456 pp. 288-337. 123456 123456

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123456 Adaptive Radiation in Hawaiian 123456 123456

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Honeycreepers: Questions on the 123456

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Reading 123456

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1) Define adaptive radiation, and explain its relationship to endemic species. Give one example of 123456

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adaptive radiation in Hawaiian species other than honeycreepers. 123456

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123456 2) Why are fossil records valuable sources of information to scientists studying the evolution of 123456 123456

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native bird species? 123456

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123456 3) Is the shift in the size of ÿiÿiwi bills over the last 100 years an example of adaptive radiation in 123456 123456

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action? Explain your answer. 123456

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123456 4) Using what you have learned about evolution and adaptation, explain why extinctions of rain 123456 123456

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forest bird species have happenedand continue to occurso rapidly in the face of human-caused 123456

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changes to native Hawaiian rain forests. 123456

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