This course will stretch your thinking about evolutionary biology by considering practical elements of defining species. We will consider diverse concepts and definitions, and focus on the utility of each for biodiversity, management, and practical discussion in evolutionary biology and ecology.
This isn't just a question about beetles, or butterflies, or barnacles. What is the meaning of distinct diversity among pathogens, how does it influence our understanding that there are Drosophila other than melanogaster? Did you know that on average over 1200 species have been described every year since Linnaeus (1758)? Why get excited when a new whale species is described -- in 2021?
You'll be expected to READ primary literature and reviews, DISCUSS complex questions of biodiversity, and WRITE a rolling synthesis of your understanding of this question: what is a species?
We meet Wednesdays 1020am, C130 Life Sciences Building
Jan. 11 -- [ s l i d e s ]
Today we discuss informally the origin of species: that living things are distinct. We take it for granted, but the history of the concept and the utility to conservation/management scientists as well as indigenous people outside of western science are both important ways of understanding biodiversity. We will have an interactive discussion of how you think about species diversity on a regular basis, how you use this information and refer to species that we see every day -- focusing on species that did NOT have a genome a decade or more ago, but those that are part of our lives. How many recognized species in the same genus can you name that can be found around here?
For next week, there are a lot of options we could start with, but let's start on typical GENE territory of thinking about species distinctions and origins by reading Grant & Grant 2009 .
Jan. 18 -- [ s l i d e s ]
The purpose of "species" is important for communication and reference, for the measurement of biodiversity. So, it has an extrinsic meaning beyond the definition that we need to understand. So we will get into the utility of species, how the term is used for management, conservation, ecology, biodiversity - and yes for the study of speciation - because we are talking about what is separable, and in evolutionary terms separation involves time.
for next week response paper no. 1: "Given my background and what we have just discussed (Grant & Grant 2009), what does it mean to be a species?" For information on expectations see Grading. Due at beginning of class. For all papers, please PRINT your paper, INCLUDE your name, and PAY ATTENTION to the rubric.
for next week we will read an example of how biologists go about DEFINING new species. We visit the UGA Herbarium next week, and our exploration and discussion there will require this background.
Jan. 25 -- visit to UGA Herbarium (Miller Plant Sciences, 2nd floor, room 2501) to see collection tour and type specimens.
I will return my evaluation of response paper no.1 by next week so you can calibrate your effort for remaining assignments.
for next week we will read TWO papers, Harpole & Tilman to remind ourselves that species DO different things, and also read why tree species are important! (really, why the fact there are distinct species is important) yes, this is your Ecology Immersion Week... :)
Feb. 1 -- [ s l i d e s ]
So today we will discuss some of the background on the fact that species being distinct from one another is ecologically important. I know you had two papers, but given the scope of the class you can look at the Resources and see there are so many other directions to go ranging from the experimental to the theoretical; I've tried to keep it straightforward for this class.
Where we will go next is to talk more directly about how species form? This will at first involve the Biological Species Concept and Dobzhansky-Muller interactions, but are these properties of species? Or of gene regions?
for next week response paper no. 2: "Given my background and what we have just discussed, what does it mean to be a species?" For information on expectations see Grading. Due at beginning of class. For all papers, please PRINT your paper, INCLUDE your name, and PAY ATTENTION to the rubric. Yes, the assignment is the same. But now you have seen and read more. Build on that information. So, reconsider and increase the complexity, nuance, or other elements of your response. It is OK to build on the information etc. from previous papers; you are learning how to refine your communication of complex ideas.
Reading for next week asks, "what is reproductive isolation?"
Feb. 8 -- [ s l i d e s ]
Now we are getting into the heart of what is often evaluated in evolutionary biology with respect to speciation. We will discuss the paper by Westram et al., which is pretty information-dense. Remember our goal is to keep updating our understanding of the process of speciation as well as the reality of the entities we call species!
for next week response paper no. 3: "Given my background and what we have just discussed, what does it mean to be a species?" For information on expectations see Grading. Due at beginning of class. For all papers, please PRINT your paper, INCLUDE your name, and PAY ATTENTION to the rubric.
Reading for next week is a response paper to Westram et al., published in the same issue, by Dr. Leonie Moyle.
Feb. 15 -- [ s l i d e s ]
OK, so you maybe getting saturated with the idea of reproductive isolation, but we will continue our discussion of the idea today and then get into an instance where this type of research is being applied with results that are a bit of a conundrum.
for next week response paper no. 4: "Given my background and what we have just discussed, what does it mean to be a species?" For information on expectations see Grading. Due at beginning of class. For all papers, please PRINT your paper, INCLUDE your name, and PAY ATTENTION to the rubric.
Reading for next week is a preprint study on two species of the flowering plant Mimulus.
Feb. 22 -- [ s l i d e s ]
Now let's mix it up a bit, we are going to finish up our discussion of reproductive isolation that focuses on intrinsic e.g. genomic mechanisms and include a recent paper highlighting work that evaluates the role of sexual selection on speciation.
for next week response paper no. 5: "Given my background and what we have just discussed, what does it mean to be a species?" For information on expectations see Grading. Due at beginning of class. For all papers, please PRINT your paper, INCLUDE your name, and PAY ATTENTION to the rubric.
For next week lets read about sexual selection as a component of speciation: Mendelson & Safran 2021
Mar. 1 -- [ s l i d e s ]
Our last class before spring break, we wrap up discussing sexual selection.
for next week (week after spring break) response paper no. 6: "Given my background and what we have just discussed, what does it mean to be a species?" For information on expectations see Grading. Due at beginning of class. For all papers, please PRINT your paper, INCLUDE your name, and PAY ATTENTION to the rubric.
for our class session on March 15, we will read and prepare to discuss a nice meta-analysis of how pre-mating and postmating isolation seem to be involved in recent studies.
March 6-10 University of Georgia Spring Break!
Mar. 15 -- [ s l i d e s ] Where we are going to start going is talking more about the component of time involved in speciation. This unites our background on drift, selection, mutation, isolation, assortative mating, and all the ways that frequencies of particular elements of genomic diversity change through time. We will discuss today the paper you read over Spring Break. Or, really, in the last 48 hours is more likely, which is fine.
for next week response paper no. 7: "Given my background and what we have just discussed, what does it mean to be a species?" For information on expectations see Grading. Due at beginning of class. For all papers, please PRINT your paper, INCLUDE your name, and PAY ATTENTION to the rubric.
for next week, we will read a CLASSIC paper by Kevin de Queiroz.
Mar. 22 -- [ s l i d e s ]
Okay, so obviously we have to learn how to think about a whole diversity of ways of defining biodiversity because most organisms we simply cannot evaluate reproductive isolation, or behavioral interactions. We just know where they are, that they look distinct, maybe that we have genomic information relative to other similar species.
for next week response paper no. 8: "Given my background and what we have just discussed, what does it mean to be a species?" For information on expectations see Grading. Due at beginning of class. For all papers, please PRINT your paper, INCLUDE your name, and PAY ATTENTION to the rubric.
for next week, we will read a kind-of-hairy paper (Hudson & Coyne 2003) on where genealogical divergence and reproductive isolation might find a common ground...
Mar. 29 -- [ s l i d e s ]
yikes, that is a tough one but we have to know that people are grounding their understanding of overall divergence into the mechanisms of organisms becoming distinct in a way we can all agree upon. We will discuss and talk about how simulation is used in evolutionary biology.
for next week (week after ACC break) response paper no. 9: "Given my background and what we have just discussed, what does it mean to be a species?" For information on expectations see Grading. Due at beginning of class. For all papers, please PRINT your paper, INCLUDE your name, and PAY ATTENTION to the rubric.
for next week (actually, we will read a paper that asks whether divergence patterns alone are enough to consider whether or not two populations of organisms represent distinct species, by my old friend Mike Hickerson.
April 3-7 ACC public schools spring break, you catch a break as well...this week we interrupt our regularly scheduled species discussion for a GENE program assessment, Dr. Rodney Mauricio will guide the class this day to discuss the GENE curriculum :)
April 12
Today we will discuss one more paper (Hickerson et al) that deals with quantifiable divergence among samples of biodiversity, and recognizes that even these 'objective' mechanisms have cases which will mislead us about what a species "is".
This, along with the Hudson & Coyne paper we last discussed, is really about time as a definition of speciation, in a sense. Genomic divergence follows predictable patterns with time, and is that enough to help us know how to define species?
So, we are near the end of the semester. And a scheduling quirk means that last week's GENE assessment will instead be NEXT week, April 19. But I have two more ways I want you to think about this problem, and some of you will need to add response papers to both to match our revision/return goals on the "What does it mean to be a species?" writing you are doing.
Keep reading please...
That means this is your last assignment from me, but it is a bigger one!
First, I want to address the whole domains and beyond of life (?) that we haven't discussed, how do any of these species concepts apply to asexual/unicellular/bacterial/viral diversity? So, two short papers:
One that is an overview of the proposed method for handling some elements of naming such diversity. (Note, these are the hyperlinks for these paper!)
You can treat these as your assigned reading to respond to by next week.
Second, I want us to deal with how decisions are made with respect to conservation or management, are made by funding decisions from both public and private agencies. How are variants within species recognized?
We will read: https://onlinelibrary.wiley.com/doi/full/10.1046/j.0962-1083.2001.01411.x because it synthesizes across a big end-of-last-century argument, but I was tempted to assign other parts of that argument (https://conbio.onlinelibrary.wiley.com/doi/10.1046/j.1523-1739.1997.96109.x ) or even better, looking at this problem from the perspective of legal definitions of protected organisms (Holly Doremus, 1997). Those other two links are purely optional, but might be your thing.
Your last paper(s) includes these last TWO assigned readings (the microbial species naming paper, and Fraser & Bernatchez 2002) as a component. This last paper (same assignment as always) is required to be one of the papers you turn in, i.e. it will be counted as one of the 8 from which your grade is determined.
for TWO WEEKS FROM NOW response paper no. 10: "Given my background and what we have just discussed, what does it mean to be a species?" For information on expectations see Grading. Due at beginning of class. For all papers, please PRINT your paper, INCLUDE your name, and PAY ATTENTION to the rubric.
April 19 Dr. Rodney Mauricio will guide the class this day to discuss the GENE curriculum
Next week we will turn in our last reading/discussion synthesis revisions. You should be starting to feel pretty comfortable with what things you feel are most important in making these important decisions about how biodiversity is named. We won't discuss the papers this week, but will spend time on both of the recent readings as well as overall synthesis.
April 26 [ s l i d e s ]
OUR GRAND SYNTHESIS
Every component of grading will follow the EMRN rubric, effectively a 4-level system for identifying how I perceive you showing your understanding. The rationale for this approach is explained here. The rubric applies to your communication/writing quality, scholarship (and proper citation thereof), and the specificity/detail/thoughtfulness of your response.
Grades will be assigned to a series of eight response papers that will be described; more may be assigned but only your top eight count towards your grade. If you receive an R or an N, you may turn in a revised assignment up through the last day of classes, no later. The assignment will be the same each time it comes up: "Given my background of understanding and what we have just discussed, what does it mean to be a species?" Each assignment is expected to be one page, as densely packed as you can make it with your insights (so, yes, single spaced). There must be citations of the resources you use (in particular the most recent ones), but otherwise creativity counts -- using figures, illustration, comics, songs, poetry are certainly ways to present your engagement!
Obviously, at the end of the semester I'm required to translate these assessments into traditional grades. It is my hope that being transparent about the rationale for grading makes it easy for you to target an easily defensible "good" grade.
Doing well (better than "Meets Expectation") requires your engagement in the class – which includes preparation for class, focus during our activities, presence and responsiveness, asking questions by whatever format, listening to others, referring to specific ideas from readings/discussion, and synthesis of all this information in your reports.
Additionally, it is required that you complete the three assessments from the Department of Genetics as part of your capstone 4950 experience.
This is just a simple one-page schedule (for a 1-credit class!) and advertisement to GENE and other interested folks, goal is not to grade intensely but foster discussion, practical examples, reading, and keeping an eye out for congeners that we see every day.
As interesting links, additional papers, or other resources come up they will be posted here. There is no way y'all can read everything cool that I can think of in the allotted time, but some may seem of particular interest and will guide your written inquiry into "what does it mean to be a species?"
Papers considered but not chosen for weeks 1, 2: Mark Vellend's work equating population genetic theory into community ecology models, evaluations of the Neutral Theory of Biodiversity, an earlier paper by David Tilman on species diversity itself providing ecosystem benefit, and a really cool manipulative experiment testing the "neutral" function of species in a community
Papers considered but not chosen for late March: Schluter & Riesenberg: https://www.pnas.org/doi/10.1073/pnas.2122153119 comes after Hickerson. Also, obviously there are folks who are thinking very clearly about how to delimit species even when reciprocal monophyly is not achieved by the markers we are looking at, Lacey Knowles and Bryan Carstens are pretty brilliant colleagues and this is a useful paper to consider.
As of 9-25-22, the assigned authorship list includes among the First/Senior authors of each reading: men 17, women 7, unknown 1. BIPOC count unknown. How can I find articles for us to read that represent the balance of human diversity, even in science, better? If students come across a good representative article on modes of species definition that helps illuminate the many people contributing to this field, Dr. Wares will appreciate being alerted!
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