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Author Topic:   Definition of Species
RAZD
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Message 1 of 450 (420824)
09-09-2007 5:26 PM


I think everyone is fairly familiar with the biological species definition, and some may be familiar with the morphological definition. Here is a site for reference on these definitions:

U of Michigan Lectures - The Process of Speciation

quote:
  • Biological species concept: This concept states that "a species is a group of actually or potentially interbreeding individuals who are reproductively isolated from other such groups."

  • Morphological species concept: Oak trees look like oak trees, tigers look like tigers. Morphology refers to the form and structure of an organism or any of its parts. The morphological species concept supports the widely held view that "members of a species are individuals that look similar to one another." This school of thought was the basis for Linneaus' original classification, which is still broadly accepted and applicable today.
  • Where we can study living populations of sexual species we can use the first definition, but when we deal with the fossil record or with asexual species we would have to use the second definition.

    There is also another definition in the forum glossary:

    http:///WebPages/Glossary.html#S

    quote:
    A basic taxonomic category for which there are various definitions. Among these are an interbreeding or potentially interbreeding group of populations reproductively isolated from other groups (the biological species concept) and a lineage evolving separately from others with its own unitary evolutionary role and tendencies (Simpson's evolutionary species concept). Employing the terms of population genetics, some definitions can be combined into the concept that a species is a population of individuals bearing distinctive genes and gene frequencies, separated from other species by biological barriers preventing gene exchange.

    Buried in Message 118 is this comment:

    The whole field of classification began with Morphology. The biological definition was first proposed by Ernst Mayer in 1942. Scientists are still attempting to find better ways to distinguish species using genetics (I'll talk about that later). In any case, while the biological definition is nice on paper it is not as commonly used in practice as indicated by this university:
    http://www-geology.ucdavis.edu/...ofLife/speciesconcept.html

    To quote "However, in the real world, it is time-consuming and expensive to make the observations of organisms in their real habitat that would allow us to say with confidence that such-and-such a set of organisms really is a species. And in the fossil world, it is impossible. So instead, most biologists and all paleontologists make a good-faith guess about the boundaries of the set of organisms they propose to name a species. Typically, the species is defined on the morphology it has, not on the genetics and behavior that is specified in the biological species concept. ... "

    The news article listed by Murkywater above has a link to the actual Journal article:

    http://www.allenpress.com/pdf/mamm-87-04-24_643..662.pdf

    quote:
    We define a genetic species as a group of genetically compatible interbreeding natural populations that is genetically isolated from other such groups. This focus on genetic isolation rather than reproductive isolation distinguishes the Genetic Species Concept from the Biological Species Concept. Recognition of species that are genetically isolated (but not reproductively isolated) results in an enhanced understanding of biodiversity and the nature of speciation as well as speciation-based issues and evolution of mammals. We review criteria and methods for recognizing species of mammals and explore a theoretical scenario, the Bateson-Dobzhansky- Muller (BDM) model, for understanding and predicting genetic diversity and speciation in mammals. If the BDM model is operating in mammals, then genetically defined phylogroups would be predicted to occur within species defined by morphology, and phylogroups experiencing stabilizing selection will evolve genetic isolation without concomitant morphological diversification. Such species will be undetectable using classical skin and skull morphology (Morphological Species Concept).

    -- Journal of Mammalogy, 87(4):643-662, 2006 p.643 (abstract)


    Now my first impression is that this is really just the biological definition of species using genetics to determine reproductive isolation, and one that would be useful for finding "cryptic" species, one that could be applied to asexual species, and even extended to some fossils (where DNA is recoverable).

    quote:
    We define genetic species as a group of genetically compatible interbreeding natural populations that is genetically isolated from other such groups. Under our definition of the Genetic Species Concept, speciation is the accumulation of genetic changes in 2 lineages (Bateson 1909) that produce genetic isolation and protection of the integrity of the 2 respective gene pools that have independent evolutionary fates. Therefore, the process of speciation depends on divergence in genes, the genome, and chromosome structure (Check 2005), and species exist when the integrity of 2 gene pools is protected as a consequence of genetic differences in their respective genomes (e.g., as outlined in the Bateson-Dobzhansky-Muller [BDM] model but not restricted to those conditions).

    -- ibid p.645


    Reading further it seems that they establish a somewhat arbitrary delineation for "type" species concept based on <0.5% difference in the mitochondrial cytochrome-b gene:

    quote:
    Sister species of mammals that have been recognized as species based on morphology often have cytochrome-b distance values .5% and this magnitude of divergence in the cytochrome-b gene has been associated with taxa recognized as species (Bradley and Baker 2001).

    -- ibid p.649


    I would think you really want to compare the total genome to ensure you are picking up where the genetic change is occurring within the population(s). I also think you would want to do a statistical mapping of all the variations by frequency and see if you have one or more peaks in the data, and let those peaks define your species (or incipient species depending on the degree of overlap and geographical separation).

    Furthermore, where you draw the line (>0.5%) could determine arbitrary species designations, with several monophylic species with wide viable hybrid zones, as opposed to one polyphylic species, being a matter of somewhat subjective interpretation.

    quote:
    Genetic isolation is not simply an off-and-on switch because genetic changes in allopatric populations are accumulated slowly across the genome and may involve a substantial number (estimated at 200 for Drosophila—Presgraves 2003) of loci affecting isolation. Accumulation of adequate change in independent sister lineages that results in genetic isolation will be a chance event occurring rapidly in some cases but requiring long periods of separation in other cases. Genetic isolation resulting from the BDM model will be expected to produce intermediate and incomplete stages of reproductive isolation before the completion of reproductive isolation. We predict that genetic profiles of interactions between members of mammalian phylogroups will reveal examples of complete genetic isolation and examples of essentially no genetic isolation even with the same genetic divergence in the mitochondrial marker used to select phylogroups for more intensive study. But, more commonly in phylogroups with .5% genetic distance in the cytochrome-b gene, various combinations of genetic isolation will be apparent. As a result, when phylogroups are sympatric, there will often be hybridization, and data documenting the genetic basis for any level of isolation will be difficult to organize into well-defined stages (Table 2). Genetically defined hybrid zones will be common.

    -- ibid p.651


    Now it is not surprising to me that there would be grades of separation to be found in the data, as various populations would be at different points in developing isolating mechanisms to achieve speciation, as this is a gradual process after all.

    Nor am I surprised to think that there could be subpopulations that don't have the same proportions of alleles as other subpopulations, due to geographic factors that would make direct mating difficult, and where gene flow would lag temporally.

    The question though is whether any member of subpopulation (A) could mate with a member of subpopulation (B) and produce viable (hybrid) offspring ... and this is not really answered when you have the subpopulations defined by average genetic similarity around mean values (there will always be a distribution in any population) and only observe mating between some of the two populations (that may or may not be statistically high in either subpopulation). You don't know who is in the "hybrid zone" when mating occurs.

    Why is the definition of species important and what is the use for the definition of species?

    Speciation is is the dividing line between what are considered microevolutionary and macroevolutionary processes and mechanisms, between the generation of homogeneous change within a population (evolution), and the generation of heterogeneous change (diversification) between diverging (especially for new) species.

    Thus I would define any population with a single peak frequency distribution as a species, any population with two peaks and a high "saddle" between them as incipient species, and any population with two peaks and a low "saddle" as different species. Analysis of this type of pattern for species like horses, zebras and donkeys would give you an idea of the saddle height necessary for speciation.

    Comments?

    Enjoy.

    Edited by RAZD, : transfer flow

    Edited by RAZD, : fixed symbols, signature


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    Replies to this message:
     Message 3 by Doddy, posted 09-10-2007 5:07 AM RAZD has responded
     Message 5 by Ben!, posted 09-10-2007 10:55 AM RAZD has responded
     Message 20 by RAZD, posted 11-14-2007 3:07 PM RAZD has responded
     Message 22 by RAZD, posted 06-19-2009 8:34 AM RAZD has acknowledged this reply
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    AdminNosy
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    Message 2 of 450 (420831)
    09-09-2007 5:41 PM


    Thread moved here from the Proposed New Topics forum.
      
    Doddy
    Member (Idle past 4019 days)
    Posts: 563
    From: Brisbane, Australia
    Joined: 01-04-2007


    Message 3 of 450 (420897)
    09-10-2007 5:07 AM
    Reply to: Message 1 by RAZD
    09-09-2007 5:26 PM


    Why is the definition of species important and what is the use for the definition of species?

    I don't think it is important. It will always drawing an arbitrary line on a gradual process, and I don't think it even matters.

    Speciation is is the dividing line between what are considered microevolutionary and macroevolutionary processes

    Micro and macro evolution, being defined as speciation and morphological changes, are dependant on species as a unit (at least in biology, rather than creation science). But species aren't units, and so dividing micro from macro is just as problematic and useless as diving species from each other. It just leads to the question of 'why is macroevolution an important term?'.

    In conclusion, a one-size fits all method of determining species will always cause problems. It's subjective, and thus useless outside of the obviously distinct species.

    That's my opinion anyway.


    Help to inform the public - contribute to the EvoWiki today!

    What do you mean "You can't prove a negative"? Have you searched the whole universe for proofs of a negative statement? No? How do you know that they don't exist then?!


    This message is a reply to:
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    RAZD
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    Message 4 of 450 (420914)
    09-10-2007 8:42 AM
    Reply to: Message 3 by Doddy
    09-10-2007 5:07 AM


    But species aren't units, and so dividing micro from macro is just as problematic and useless as diving species from each other.

    It's two aspects of the same issue. But speciation - the division of one species into two or more species - is what accounts for the diversity in types of organisms, and having a usable definition of species lets you determine when speciation has occurred, and then you can study how long it takes and what the specific mechanisms are.

    In conclusion, a one-size fits all method of determining species will always cause problems. It's subjective, and thus useless outside of the obviously distinct species.

    Yet we know that many populations of organisms are genetically isolated by lack of mechanism for sharing genes (sex or horizontal transfer), and that as a result they have different resources to use when reacting to changing ecological situations or opportunities.

    The concept of species allows better understanding of the mechanisms that lead to (a) continual change in a genetic line of a population of otherwise similar organisms or (b) diversification of life into different ecological opportunities.

    Enjoy.


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    This message is a reply to:
     Message 3 by Doddy, posted 09-10-2007 5:07 AM Doddy has responded

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    Ben!
    Member (Idle past 1731 days)
    Posts: 1154
    From: San Diego, CA
    Joined: 10-14-2004


    Message 5 of 450 (420930)
    09-10-2007 10:55 AM
    Reply to: Message 1 by RAZD
    09-09-2007 5:26 PM


    Now my first impression is that this is really just the biological definition of species using genetics to determine reproductive isolation, and one that would be useful for finding "cryptic" species, one that could be applied to asexual species, and even extended to some fossils (where DNA is recoverable).

    I wouldn't phrase it that way, but reading purely what's in your post, it seems the same to me.

    Reading further it seems that they establish a somewhat arbitrary delineation for "type" species concept based on <0.5% difference in the mitochondrial cytochrome-b gene:

    The number seems arbitrary as they haven't defined what "often" means. Also seems that it's lacking any theoretical underpinning, and is pulled straight from the data with no attempt at providing additional theoretical "motivation" for choosing the number.

    Thus I would define any population with a single peak frequency distribution as a species, any population with two peaks and a high "saddle" between them as incipient species, and any population with two peaks and a low "saddle" as different species. Analysis of this type of pattern for species like horses, zebras and donkeys would give you an idea of the saddle height necessary for speciation.

    Razd, sorry, I didn't understand what frequency distribution you were talking about here, or in the other part of your post. Could you elaborate?

    Sorry I couldn't come up with any better comments... but thanks for the information.


    This message is a reply to:
     Message 1 by RAZD, posted 09-09-2007 5:26 PM RAZD has responded

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    RAZD
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    Message 6 of 450 (420942)
    09-10-2007 11:42 AM
    Reply to: Message 5 by Ben!
    09-10-2007 10:55 AM


    Ben, long time no cross paths.

    Razd, sorry, I didn't understand what frequency distribution you were talking about here, or in the other part of your post. Could you elaborate?

    Well, every individual would be genetically unique, so there is no genetic "type" in a strict sense, but a distribution. You would need some way to map this distribution that wouldn't bias the data, such as take a sample from a sister species that you know is isolated reproductively and genetically, and then count the differences between each individual in the population and that sister species sample. It should show a frequency distribution like a bell curve if the population is relatively homogeneous for genotype (one species), but it should have multiple peaks if it is heterogeneous for genotype (two or more species or incipient species).

    Because of gene mixing (reproductive imperfection) you probably could not make a cladogram for the population using the whole genomes, as there would be cross branching (some genes from one parent line others from the other at every level).

    The number seems arbitrary as they haven't defined what "often" means. Also seems that it's lacking any theoretical underpinning, and is pulled straight from the data with no attempt at providing additional theoretical "motivation" for choosing the number.

    That's kind of my impression: more work needed to refine the application.

    Enjoy.


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    Doddy
    Member (Idle past 4019 days)
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    Message 7 of 450 (421024)
    09-10-2007 7:26 PM
    Reply to: Message 4 by RAZD
    09-10-2007 8:42 AM


    RAZD writes:

    It's two aspects of the same issue. But speciation - the division of one species into two or more species - is what accounts for the diversity in types of organisms

    Branching is what accounts for the structure of the tree, but it is completely useless to try to distinguish the exact line between a bough, a branch and a twig. You don't really need to know what the precise definition of a twig is to study how they form.

    RAZD writes:

    having a usable definition of species lets you determine when speciation has occurred, and then you can study how long it takes and what the specific mechanisms are.


    So, we need to have a definition for species to study how species form. Well, that just brings me to the questions of: why do we want to know exactly when speciation has occured? We don't really need to know that to understand the processes involved.


    Help to inform the public - contribute to the EvoWiki today!

    What do you mean "You can't prove a negative"? Have you searched the whole universe for proofs of a negative statement? No? How do you know that they don't exist then?!


    This message is a reply to:
     Message 4 by RAZD, posted 09-10-2007 8:42 AM RAZD has responded

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    RAZD
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    Message 8 of 450 (421048)
    09-10-2007 9:26 PM
    Reply to: Message 7 by Doddy
    09-10-2007 7:26 PM


    Well, that just brings me to the questions of: why do we want to know exactly when speciation has occured?

    Well even if you don't I do. Understanding this process can help understand why some speciations are (relatively) abrupt and morphologically distinct while othes are cryptic.

    Understanding the cryptic species in mosquitos was important to distinguish those that were carriers of malaria and those that were not, when they have all been considered to be one species before.

    Enjoy.


    Join the effort to unravel AIDS/HIV, unfold Proteomes, fight Cancer,
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    we are limited in our ability to understand
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    RebelAAmericanOZen[Deist
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    Medis
    Member (Idle past 2000 days)
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    Message 9 of 450 (429146)
    10-18-2007 5:09 PM


    Complexity of an organism
    Not sure if this is the right place to post this but...

    How do you define the complexity of an organism?

    One evidence of the occurrence of macroevolution is the fossil record observation that invertebrates were on the earth much longer than vertebrates. Thus showing that "complex" organisms evolved from "simple" organisms. But how do you define a complex organism? Having more cells? Having a skeletal structure? Is it possible to say that an organism with a skeletal structure is "more complex" than an organism without it and why?

    Edited by Malangyar, : No reason given.

    Edited by Malangyar, : No reason given.


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    Dr Adequate
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    Message 10 of 450 (429162)
    10-18-2007 5:31 PM
    Reply to: Message 9 by Medis
    10-18-2007 5:09 PM


    Re: Complexity of an organism
    How do you define the complexity of an organism?

    We don't.

    Is it possible to say that an organism with a skeletal structure is "more complex" than an organism without it ...

    No, it isn't.

    Well, to be precise, which I am even in my sleep, it is possible to say it, but I challenge anyone to mean anything by it.

    Edited by Dr Adequate, : No reason given.


    This message is a reply to:
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    Medis
    Member (Idle past 2000 days)
    Posts: 34
    Joined: 10-16-2007


    Message 11 of 450 (429169)
    10-18-2007 5:48 PM
    Reply to: Message 10 by Dr Adequate
    10-18-2007 5:31 PM


    Re: Complexity of an organism
    Dr Adequate writes:

    No, it isn't.

    Well, to be precise, which I am even in my sleep, it is possible to say it, but I challenge anyone to mean anything by it.

    Are you sure this is true, I mean I'm looking at a wikipedia article right now saying "More complex organisms such as...". I mean can't you say that a multicellular organism is more complex than a single-celled organism?


    This message is a reply to:
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    Chiroptera
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    Message 12 of 450 (429172)
    10-18-2007 5:58 PM
    Reply to: Message 11 by Medis
    10-18-2007 5:48 PM


    Re: Complexity of an organism
    I mean can't you say that a multicellular organism is more complex than a single-celled organism?

    Sure, you can say it. But what does it mean?

    Edited by Chiroptera, : No reason given.


    In many respects, the Bible was the world's first Wikipedia article. -- Doug Brown (quoted by Carlin Romano in The Chronicle Review)
    This message is a reply to:
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    Medis
    Member (Idle past 2000 days)
    Posts: 34
    Joined: 10-16-2007


    Message 13 of 450 (429175)
    10-18-2007 6:01 PM
    Reply to: Message 12 by Chiroptera
    10-18-2007 5:58 PM


    Re: Complexity of an organism
    That it has more parts and thus has a more interconnected whole than an organism with fewer cells.
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    Dr Adequate
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    Message 14 of 450 (429182)
    10-18-2007 6:36 PM
    Reply to: Message 11 by Medis
    10-18-2007 5:48 PM


    Re: Complexity of an organism
    Are you sure this is true ...

    Yes. I was taking you up on a particular example, and I challenge you, anyone else, the whole world, to demonstrate that vertebrates, in general, are more "complex" than invertebrates in any meaningful sense whatsoever.

    I mean can't you say that a multicellular organism is more complex than a single-celled organism?

    Not without knowing more about them.

    But the whole idea of "complexity" is that complex organism, the "red herring". It's not an interesting concept within evolutionary thought.

    Edited by Dr Adequate, : No reason given.

    Edited by Dr Adequate, : No reason given.

    Edited by Dr Adequate, : No reason given.


    This message is a reply to:
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    Medis
    Member (Idle past 2000 days)
    Posts: 34
    Joined: 10-16-2007


    Message 15 of 450 (429185)
    10-18-2007 6:47 PM
    Reply to: Message 14 by Dr Adequate
    10-18-2007 6:36 PM


    Re: Complexity of an organism
    So what should you use instead? That organisms aren't more complex than others, just different from others?
    This message is a reply to:
     Message 14 by Dr Adequate, posted 10-18-2007 6:36 PM Dr Adequate has responded

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