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General Ecology- UNH

Instructors have assigned the following study questions to help students grasp particular topics.

TABLE OF CONTENTS:

Adaptation

Alien invasions

Coevolution

Deformed frogs

Development and evolution

Global Warming

Human impact

Natural selection

Neutral theory

Pest control

Insect resistant crops

Population explosion

Phylogeny

Punctuated equilibrium

Species concept

Defining species

Adaptation

J.B. Losos, D.J. Irschick, and T.W. Schoener. 1994. Adaptation and constraint in the
evolution of specialization of Bahamian Anolis lizards. Evolution 48(6):1786-1798.
J.B. Losos, K.I. Warheit, and T.W. Schoener. 1997. Adaptive differentiation following
experimental island colonization in Anolis lizards. Nature 387:70-72.
T.J.Case. 1997. Natural Selection out on a limb. Nature 387:15-16.

I'd like to use these papers as a case study for natural selection and adaptation and constraint. There are 2 separate experiments presented by Losos. The 1994 paper is an example where they had predictions of evolutionary response in 2 species of lizards, but did not see any change over time. The 1997 Nature paper presents the results of a separate long term experiment using one of the species of Anolis lizards, where they claim to see adaptive evolution. The Case paper is a review of the 1997 Losos study.

Think about the criteria for evolution via natural selection. Have they demonstrated evolution? Have they demonstrated evolution via natural selection? If so, what are the strongest pieces of evidence? If not, are there pieces of information that are missing?

In the 1997 paper they use several multivariate statistics to answer the question: Are the island colonists morphologically different from the source population? They measured several morphological characters and want to determine whether the overall shape of the critters is different from different populations. They used principle component analysis to do this. In fig. la, think of the axis as two measures of shape, regardless of size (they have statistically removed the effect of size already). The x axis (PCA 1 principle component axis 1) could be thought of as lamellae width and hindlimb length. The y axis Labeled PCA 2) is essentially mass of the critter. Fig lb is another pictorial representation of the same data.

Are the experimental island and the source population morphologically distinct? In
which characters? What was the direction of the change?

The Adaptationist program. A case study: The Aquatic Ape debate.

After our general discussions about adaptation we will debate the merits of a specific case study.
On Monday in class we will choose sides for an adaptationist debate centering around the Aquatic
Ape Theory advocated by Morgan. One side will defend the Aquatic Ape theory and the
Adaptationist Program in all its glory and spender, while the other side will argue the Aquatic Ape
program is lame. Everyone is expected to participate, so please be prepared to defend your side.
** are required readings for everyone. The others are optional for each side's reference.

PRO:
Morgan, E. 1991 "Why a new theory is needed." In The Aquatic ape-Fact or Fiction.
Souvenir Press. pp. 9-22.
Mayr, E. 1988. "How to carry out the adaptationist program?" In To-Towards a new philosophy. Harvard Press. pp. 148-157.
Morgan, E. 1994. The aquatic ape hypothesis. The New Scientist 12 April. pp. 11-13.

CON:
Turner, A. 1991 "Adaptation and the Aquatic Ape." In The Aquatic ape: Fact or Fiction.
Souvenir Press. Pp. 133-141.
Reynolds V. 1991. Cold and Watery? Hot and Dusty? Our Ancestral Environment and Our
Ancestors Themselves: An Overview-" In The Aquatic Ape: Fact or Fiction. Souvenir
Press. pp. 331--341.


Coevolution

Williams and Gilbert. 1981. Insects as selective agents on plant vegetative
morphology: egg mimicry reduces egg laying by butterflies. Science 212:467-469.
Gilbert. 1982. The coevolution of a butterfly and a vine. Sci. Am. 247:110-121.

On Thursday we have the rare opportunity to talk with one of the authors of the papers we are reading. Dr. Larry Gilbert from the University of Texas, Austin will be visiting Cornell and has agreed to meet with this class. Texas has one of the best programs in evolutionary biology in the world, and Larry has been a leader in the department. For over 30 years, he has studied the evolution and ecology of Heliconius butterflies and Passiflora vines in the New World Tropics, especially in Costa Rica. He has also been responsible for helping set up and manage one of Costa Rica's National Parks. More recently, he has worked on the biological control of fire ants which destroy the native insect communities of the southern U.S., besides being a major pest of humans and livestock. His students have worked on a bewildering array of topics in evolutionary biology and on a range of organisms from insects to amphibians to mammals.
You should read the above papers carefully, answer the study questions, and come to section primed to ask Larry questions about coevolution, things you didn't understand in the paper, other things you may want to know (is there heritable variation in the morphology of egg mimics? have you applied a phylogenetic test to any of your evolutionary scenarios, what were the results?). If you have other, broader questions you'd like to ask about the tropics or other topics, that's fine too, but try to keep your questions as evolutionary and relevant as possible.
1) What is coevolution (be sure to mention natural selection in your answer)?
2) What reason does Gilbert provide for why coevolution is unlikely in some systems
(like temperate oak, maple systems with their parasites) and likely in others?
3) Do you agree that the evolution of egg mimicry would be correlated with expanding into new habitats? Why should the presence or absence of egg mimics limit the habitat a vine occurs in?
4)There are a large number of evolutionary scenarios given in both of the papers (eg., the one mentioned in question 3, those having to do with leaf shape, that egg mimicry evolved recently, etc.). At the time of these papers, a phylogenetic perspective had not yet really infused itself into evolutionary biology. That didn't happen until the late '80's and early '90's.
Choose one of the scenarios and briefly discuss how you could test it with a phylogeny. Draw a phylogeny or two to show the different possibilities.

Development and evolution

Oster, G. and P. Alberch. 1992. Evolution and bifurcation of developmental programs
Evolution 36 (3):444-459.
Murray, J.D.1988. How the Leopard Gets Its Spots. Scientific American. 258 (3):80-87.

What processes explain the production of discontinuous pheotypes from continouos genotypic change? What role does development play in explaining the diversity of phenotypes we see in nature? We have spent much of the semester analyzing how natural selection creates and maintains variation in nature. But is that it? Is natural selection sufficient in explaining the diversity of life?

Oster and Alberch ask: how can developmental process control th eexpression of morphological variaiton and generate the rules of phenotypic change? They propose a model for the development of skin organs like fathers, teeth, hair, etc. Please pay specific attention to the Introduction and Part I and V where they put their model in a broad contextual background. Note Fig. 11.

Murray writes about a related model (a reaction diffusion model) of how pattern formation in mammal coats develops. This simple model simulates zebra stripes, and leapord spots without using genetics. What is a morphogen?

Global Warming

1. Where did the idea of global warming originate?
2. Who have been the major players in the scientific debate over global warming?
3. What are the facts that everybody agrees on?
4. What are the main areas of disagreement?
5. How have the answers to #3 and #4 above changed over the last 10 years?
6. What are some possible consequences of global warming?
7. What are some possible consequences of increased atmospheric carbon-dioxide levels,
other than global warming?
8. What are some of the social / economic / political barriers to decreasing carbon-dioxide
emissions?

Human impact

1. What are the basic requirements (in general) that must be met for an organism / species
to survive and reproduce in a given location?
2. How have humans altered their environments in the process of trying to meet these basic
requirements?
3. Are some human-caused environmental problems inevitable because humans are "just
doing what organisms do" in order to survive and reproduces



Natural selection

Bethell, T. 1976. Darwin's mistake. Harper's Magazine 252:70-75.
Gould, S. J. 1977. Darwin's untimely burial. pp. 39-45 in Ever Since Darwin. W. W.
Norton, New York, NY.

The Bethell article caused quite a stir when it was published, even among evolutionary biologists. It brought home an issue raised in the philosophical literature about the scientific and theoretical status of natural selection. Gould responded almost immediately, but it is still not clear today that evolutionary biologists have come to grips with the issues Bethell's article raises. The issues are actually fairly complex and will require some thought on your part. I suggest reviewing your class notes on natural selection and the handout from the natural selection exercise. The following study questions are somewhat difficult, but are designed to get you thinking.

1) Use a dictionary to define tautology. In what sense is Bethell using the word? Is there a broader sense of the word 'tautology' which might apply to the presentation of natural selection from lecture?
2) What are the issues that Bethell raises with regard to natural selection?
3) In your opinion, does Gould address the issues raised by Bethell? Why or why not?

Boag and Grant, 1981.The Beak of the Finch, Ch. 1-7;

1. What is sexual selection? How is it different from natural selection? How did it manifest
itself in the finches?
2. If you were to try and measure the fitness of a finch, what would be your best surrogate
variable?
3. Why do abiotic factors play such an important role in the life of a finch? Is the relative
importance of biotic and abiotic factors probably different or the same for mainland birds?
4. Does population size appear to be related to rates of evolution in the finches? If so, how?

Neutral theory

Kimura, M. 1979. The neutral theory of molecular evolution. Scientific American
241:98-126.

1) List 4 observations from molecular studies that Kimura thought were better explained by neutral evolution (mutation and drift) than by natural selection. Explain how each of these observations is a problem for a "panselectionist" theory of neutral evolution. That is, what would one expect to observe in these cases if selection were dominant force?
2) Briefly, explain the idea of functional constraint at the molecular level and its
relevance to the neutral theory.
3) What are some of the ways selectionists explain high levels of polymorphism at the
molecular level?
4) Evaluate Kimura's writing style and organization in the introduction and the first section, "The Evolution of Darwinism." Pay specific attention to the following: a) how well does the introduction set up the rest of the paper? Are the main issues stated and sufficiently developed? Do you feel that you know the theme of the paper after reading the first two paragraphs? b) does Kimura use strong topic sentences that set up his paragraphs? Are there any particularly good examples? any bad examples? c) Are there many stylistic flaws such as those we discussed last week - using nouns instead of verbs, adjectives or adverbs; redundancy; 'negative' rather than 'positive' statements, etc. - or is the writing clear and concise? Give examples. Are the stylistic "flaws" necessary in some cases to make the whole paper flow, or are there repeated flaws.

Pest control

1. What is a "pest"? What makes an organism a "pest"?
2. How have human modifications of the environment provided conditions that are
conducive to the development of pest problems?
3. What are the main pest control strategies that currently exist?
4. How is Integrated Pest Management (aka IPM, aka Pest Management) different from the
way that pest control has been done in the past?

Phylogeny

McDade, L. A. 1992. Pollinator relationships, biogeography, and
phylogenetics.
BioScience 42:21-26. (I will pass this paper out in class).
Morell, V. 1997. The origin of dogs: running with the wolves. Science 276:16471648. (This article gives a splendid overview of the methods and conclusions from the more technical article by Vila et al. 1997).
Vila, C. et al. 1997. Multiple and ancient origins of the domestic dog. Science
276:1687-1689 plus the letters that followed and the response to those letters.

The purpose of reading these papers is to provide an introduction to the sorts of questions that phylogenies can help answer and to how we make inferences about the past and the course of evolution from phylogenies. The McDade paper focuses on 2 questions: 1) is pollination by a particular group of hummingbirds a recently derived trait in a group of plants, or is it a more ancient trait?, and 2) what is the biogeographic history of a particular genus of plants - where did they originate and did they invade new habitats once or many times? The Vila et al. paper focuses on 3 questions: 1) what is the wild ancestor of the domestic dog?, 2) how long ago were dogs domesticated?, and 3) are dogs the result of one domestication event or multiple domestication events? Overall, you should start to see the utility of what is called "tree-thinking" in answering evolutionary questions.

Some vocabulary: McDade. 1992 - trapliner - description of a foraging strategy used by pollinators in which they proceed from one plant to another, often over great distances and often repeatedly in the same order. I think this term is derived from the fur trapping industry. Trappers would often set traps up and down the lengths of rivers (trap line) or in different areas, but would often check them in a consistent manner.

Vila et al. 1997. hal2lotyi)e - a DNA sequence that differs from homologous sequences at one or more sites (e.g. ATTGCGTTA would be a haplotype, so would ATTGCGTTC, and ATGGCGTAT, provided these sequences represented homologous places in the same gene). A haplotype is essentially an allele.

coalescence - genes have genealogies just like people do. This makes sense given that genes are one of the things passed from parents to offspring. All gene copies in a single population ultimately are descended from a single ancestral gene copy. The genealogy of genes is said to coalesce to a single common ancestor. Analogously, all of us in the classroom could theoretically trace our heritage back to a single individual from whom we are all descendants. The genealogy of individuals would coalesce at this point.

monophyletic - a taxon is monophyletic if it consists of all the known species thought to
be descended from a single common ancestral species.


1) What hypothesis was generally accepted regarding the evolution of corolla morphologies and pollination by hermit hummingbirds prior to McDade's study? Be brief.
2) How was the evolution of non-hermit pollinated species related to the invasion of new
habitats? Be brief.
3) McDade argues that obligate mutualistic relationships, like some plant-pollinator relationships, can function as constraints on evolution. That is, they could constrain the diversity of a group of plants and the habitats that the plants occur in. How could one use phylogenies to test her hypothesis of constraint? Be brief.
4) What does McDade conclude about the biogeographic history of the Aphelandra
complex she studies?

Vila et al. 1997
1 )What were Vila et al.'s main findings? Be brief.
2 )What criticisms of these findings did Federoff and Nowak raise in their letter? Be
brief.
3) How did Vila et al. respond to these criticisms? Be brief.

Punctuated Equilibrium

Gould, S. J. 1980. Is a new and general theory of evolution emerging? Paleobiology
6:119-130.
Gould, S. J., and N. Eldredge. 1993. Punctuated equilibrium comes of age. Nature
366:223-227.

Gould (1980)
1. What does Gould claim are the two major premises of the Neo-Darwinian synthesis?
2. Briefly contrast the notions of reductionism and hierarchy.
3. What is the "Goldschmidt break?"
4. What is the "Wright break?"

Gould and Eldredge (1993)
1. Briefly explain Fig. I and its significance.
2.Briefly explain the solution Futuyma proposed to the problem that although
morphological change is correlated with speciation in the fossil record, evolutionary biologists lack a convincing mechanism explaining why morphological change is centered on speciation events.

Species Concept

Real world examples to help puzzle through species concept

Mallet, J. 1995. A species definition for the Modem Synthesis. TREE 10:294-298 +
letters and replies.

Darwin titled his book On the Origin of Species...., but what are species? Defining species is one of the most contentious topics in evolutionary biology and related fields. If evolution is about the origin of species, then evolutionary biologists clearly need an unambiguous and practicable definition of species. Otherwise, how do we know what entities to study and how do we identify the entities are the purported outcome of the processes we study? Species are the things that evolve and speciate - can evolutionary theory exist without a definition of species? In discussion on Thursday we will consider the nature of "species," the qualities of various species definitions, and the implications of these definitions for studying @ 6 speciation." Following is a list of some species concepts.

Biological species concept - species are groups of actually or potentially interbreeding
natural populations that are reproductively isolated from other such groups.

Evolutionary - a species is a single lineage (an ancestral-descendant sequence) of populations or organisms that maintains its identity from other such lineages and which has its own evolutionary tendencies and historical fate.

Phylogenetic - a phylogenetic species is a cluster of organisms which is diagnosably distinct (has a unique character) and within which there is a parental pattern of ancestry and descent.

Recognition - a species is the most inclusive population of (sexually reproducing)
organisms that share a common fertilization system

Cohesion - a species is the most inclusive population of individuals having the potential
for phenotypic cohesion through intrinsic cohesion mechanisms

Ecological - a species is a lineage (or set of closely related lineages) that occupies an adadptive zone (ecological niche) minimally different from that of any other lineage in its range and which evolves separately from all lineages outside its range

Morphological (Darwin's) (see Mallet article)....... the complete absence, in a well investigated region, of varieties linking together any two closely-allied forms, is probably the most important of all the criterions of their specific distinctness."

Genotypic Cluster (Mallet) - groups that remain recognizable in sympatry because of the
morphological and genotypic gaps between them

Before Reading
1) Define the following terms: Operational, Ontological, Nominalism, Sympatric,
Allopatric
2) Give your own definition of "species" (don't look one up - just write down what you
think a species is).

During and After Reading
1)Did Darwin think of species as real entities or as constructs of the human mind?
2)Briefly describe two difficulties with Darwin's definition of species.
3)Describe difficulties with the Biological Species (Interbreeding) Concept. In
particular, describe Darwin's critique and Wallace's two points. According to Wallace, why is the interbreeding criterion for distinguishing species confused (in what way are the explanation and the thing to be explained confused)?
4)According to Mallet, why is a theory-independent definition of species needed?
5)Is the genotypic cluster definition a theory-independent definition of species?
Explain.
6) According to Mallet, why is "isolating mechanism" a useless term?
To think about :Are species units of descriptive taxonomy, units of evolution, units of both, or units of neither? Defend your answer.
(related to the above question) Are species real (do they have objective existence) or are they simply constructs of the human mind? Defend your answer.


Last updated September 26, 2001
Comments, questions or new links? Contact Anne at astork@cisunix.unh.edu