Category Archives: Evolution

Planarian: a living vessel built by unicellular organisms?

by Piter Kehoma Boll

Not long ago I talked about the peculiar genome of the freshwater planarian Schmidtea mediterranea and how it challenges our view of many cellular processes. Now what if I told you that planarians also challenge our view of what a multicellular organism is?

We all know that organisms may be either unicellular or multicellular, but sometimes it is hard to tell them apart, especially in what are called colonial organisms, in which clones of unicellular individuals may live together.

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Green algae of the genus Volvox are somehow at the boundary between unicellular and multicellular organisms. Although usually considered a colony of unicellular organisms, they behave somewhat like a multicellular organism. Photo by Frank Fox.*

So where lies the boundary between gathered unicellular organisms and true multicellular organisms? One of the ideas is that for a group of cells to be considered a single unit (an organism) they must have high levels of cooperation and low levels of conflict between each other and, perhaps more important than that, they have to be dependent on the association in order to survive.

As most recent evolutionary theories predict, cooperation increases with genetic similarity. As a result, multicellular organisms are (almost) always composed of cells having the exact same genetic material, i.e., they are all clones. We know, however, that during DNA replication mutations may occur, so that eventually at least some cells of an adult and many-celled organism may have become genetically distinct. This leads to a need to find a way to fix this problem by reincreasing genetic similarity, and the way most organisms found to do that is by allowing only one lineage of their cells, the germ cells (which produce the gametes) to generate the next generation. Thus, each transition from one generation to the next passes through a “zygotic bottleneck”, i.e., a new organism is always generated from a single original cell, the zygote, which assures that the genetic similarity is always brought back.

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A developing new egg, with an embryo that is created from a single original cell, the zygot, assuring a higher genetic similarity of the cells in the whole organism. Photo by Stéphanie Bret.**

The zygotic bottleneck is not a rule for a lot of species though. Many animals and plants are able to reproduce asexually by budding, fission or many other ways. In such cases, the new organism usually is built from several different lineages of the parent organism. For example, some succulent plants may generate a new organism from a dettached leaf and several cells of the original leaf start to reproduce and together they build the new plant and, as each lineage may have suffered different mutations, the offspring is not necessarily composed of genetically identical cells. Nevertheless, even such organisms, which are able to reproduce asexually, still retain the ability to generate zygotes through sexual reproduction, which eventually “cleans that mess”.

But in planarians things get really strange. First of all, let’s explain some basic things about planarians. They have, as you know, a remarkable regeneration ability. This happens due to the presence of stem cells called neoblasts that fill their bodies. Those neoblasts are able to generate all cell types that make up the planarian’s body. In fact, all cells in a planarian must come from neoblasts, because, as weird as it may be, all differentiated cells in a planarian body ARE UNABLE TO UNDERGO MITOSIS! Once a neoblast differentiates into any kind of cell, it is condemned to die in a few days without ever letting descendants. All cells in a planarian body are therefore constantly replaced by new ones coming from neoblasts. The only lineage of differentiated cells that is still able to reproduce is that of the germ cells, which, as in other multicellular organisms, assure that the next generation will consist of organisms with genetically homogeneous cells.

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Girardia tigrina, a freshwater planarian. Photo by Wikimedia user Slimguy.**

Several freshwater planarians, however, have lost their ability to generate sexual organs and, as a result, germ cells. In order to reproduce, they must rely on a form of asexual reproduction, which in this case happens by transversal fission of the body and posterior regeneration of the missing parts. In these populations, the zygotic bottleneck disappeared completely and, as a result, any non-lethal mutation in the neoblasts is retained in the organism, leading to a population of genetically distinct neoblasts inside a planarian.

Therefore, considering the fact that asexual planarians are not genetically homogeneous, having several different neoblast lineages in the same body, and that the neoblasts are the only cells able to reproduce and continue the species, a recent publication by Fields and Levin (see references) suggests that asexual planarians are nothing more than a very complex environment built by neoblasts in order to survive. Considering that each neoblast is an independent cell, which only needs the environment (the planarian) to survive, but does not need other neoblasts, planarians, at least the asexual ones, do not seem to have reached completely the requirements of high internal cooperation and low internal competition to be considered multicellular organisms.

We could interpret the neoblasts as unicellular organisms that live together, cooperating to build a complex environment, the planarian body, with their own sterile descendants (the differentiated cells), as if they were a group of queen ants living among sterile castes. Kind of mind blowing, huh? But it actually makes sense.

If you want to read more about it and understand every detail in this theory, read the references below.

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Maybe you’d like:

Endosperm: the pivot of the sexual conflict in flowering plants

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References:

Fields, C; Levin, M. (2018) Are Planaria Individuals? What Regenerative Biology is Telling Us About the Nature of Multicellularity. Evolutionary Biology: 1–11.

West, S. A.; Fisher, R. M.; Gardner, A.; Kiers, E. T. (2015) Major evolutionary transitions in individuality. PNAS 112 (33): 10112-10119. https://doi.org/10.1073/pnas.1421402112

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The Dying Melody: Habitat fragmentation is killing the songs of our birds

by Piter Kehoma Boll

Birds, especially passerine birds (those of the order Passeriformes) are known for their ability to produce complex and melodic calls, or songs, used for a variety of purposes. Who doesn’t love to hear the birds singing beautifully in the forest?

Passerine birds include two main groups: the Passeri (songbirds), also called Oscines, and the Tyranni (tyrants), also called Suboscines. Both groups are able to produce complex calls, but those of the Oscines are usually more smooth and melodic and sound less mechanic.

However, there is one more difference between the calls of both groups. The calls of the tyrants are genetically transmitted from the parents to the offspring, i.e., they do not need to learn how to sing with adult birds. Among the songbirds, on the other hand, the complexity of the call is largely culturally inherited, i.e., they learn how to sing with other birds of the same species.

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The orange-billed sparrow is a songbird found in the Neotropical region. Photo by Francesco Veronesi.*

A recent study published in the journal Animal Behavior (see reference) analyzed the complexity of the calls of two passerine birds found in the forests of Costa Rica: a songbird, the orange-billed sparrow Arremon aurantiirostris, and a tyrant, the scale-crested pygmy tyrant, Lophotriccus pileatus.

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The scale-crested pygmy tyrant is a tyrant found in the Neotropical region. Photo by Chris Jimenez.*

The team compared the complexity of the calls in different populations of each species living in forest fragments of different sizes. The conclusion was that, while fragment size did not affect the complexity of the call in the tyrant bird, it significantly affected the call of the songbird.

Populations of the orange-billed sparrow that live in smaller fragments show a less complex song than those living in larger fragments. As the call in this species is culturally transmitted, this reduction in complexity is most likely a result of cultural erosion. As smaller fragments only support smaller populations, the birds do not interact that much with other individuals of the same species while they are growing up, and as a result their song becomes simpler and simpler.

We, humans, are the ones to blame for that, as you may already know. Our irresponsible practices are not only reducing population size in other species, but are destroying their culture as well.

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Reference:

Hart, P. J.; Sebastián-González, E.; Tanimoto, A.; Thompson, A.; Speetjens, T.; Hopkins, M.; Atencio-Picado, M. (2018) Birdsong characteristics are related to fragment size in a neotropical forest. Animal Behavior 137: 45–52. https://doi.org/10.1016/j.anbehav.2017.12.020

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You know nothing, humans! A planarian genome challenges our understanding of how life works

by Piter Kehoma Boll

We finally have a rather complete sequencing of a planarian’s genome, more precisely, of the planarian Schmidtea mediterranea, which is an important model organism for the study of stem cells and regeneration.

In case you don’t know, planarians have a remarkable ability of regeneration, so that even tiny pieces are able to regenerate into a whole organism. They are like a real-life Wolverine, but somewhat cooler! This amazing ability is possible due to the presence of a group of stem cells called neoblasts that can differentiate into any cell type found in the planarian’s body. In fact, all differentiated cell types in planarians are unable to undergo mitosis, so that neoblasts are responsible for constantly replacing cells in every tissue. But we are not here to explain the details of planarian regeneration. We are here to talk about Schmidtea mediterranea‘s genome!

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Look at its little cock eyes saying “I will destroy everything you think you know, humans!” Photo by Alejandro Sánchez Alvarado.*

A rather complete genome of S. mediterranea has been recently published and its analysis reveal some astonishing features.

First of all, 61.7% of S. mediterranea‘s genome is formed by repeated elements. Repeated elements are basically DNA strands that occur in multiple copies throught the genome of an organism. They are thought to come from the DNA of virus that was incorporated to the host’s DNA. In humans, about 46% of the genome is formed by repeated elements. Most repeated elements of S. mediterranea belong to unidentified families of retroelements, thus suggesting that they are new undescribed families. Those repeats are very long, having more than 30 thousand base pairs, which are not known to exist in other animals. The only other group of repeated elements with a similar size is found in plants and known as OGRE (Origin G-Rich Repeated Elements). The long repeat found in Schmidtea was therefore called Burro (Big, unknown repeat rivaling Ogre).

But certainly the most surprising thing about S. mediterranea‘s genome is the lack of many highly conserved genes that are found in most eukaryotes and that were thought to be essential for cells to function properly.

Schmidtea mediterranea lacks genes responsible for repairing double-stranded breaks (DSBs) in DNA, which would make them very likely to suffer a lot of mutations and sensitive to anything that induces DSBs. However, planarians are known to have an extraordinary resistance to gamma radiation that induces DSBs. Do they have other repairing mechanisms or is our current understanding about this process flawed?

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Several “essential” genes and their presence (in green) or absence (in red) in several animals. Schmidtea mediterranea lacks them all. Image extracted from Grohme et al. (2018).**

Another important gene that was not found in S. mediterranea is the Fatty Acid Synthase (FASN) gene, which is essential for an organism to synthesize new fatty acids. Planarians therefore would have to rely on the lipids acquired from the diet. This gene is also absent in parasitic flatworms and was at first thought to be an adaptation to parasitism but since it is absent in free-living species as well, it does not seem to be the case. Could it be a synapomorphy of flatworms, i.e., a character that identifies this clade of animals?

That is not enough for little Schmidtea, though. More than that, this lovely planarian seems to lack the MAD1 and MAD2 genes, which are found in virtually all eukaryotes. Those genes are responsible for the Spindle Assembly Checkpoint (SAC), an important step during cell division that prevents the two copies of a chromosome to separate from each other before they are all connected to the spindle apparatus. This assures that the chromosomes will be evenly distributed in both daughter cells. Errors in this process lead to aneuploidy (the wrong number of chromosomes in each daughter cell), which is the cause of some genetic disorders such as the Down syndrome in humans. Planarians do not have any trouble in distributing their chromosomes properly, so what is going on? Have they developed a new way to prevent cellular chaos or, again, is our current understanding about this process flawed?

Let’s wait for the next chapters.

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Reference:

Grohme, M. A.; Schloissnig, S.; Rozanski, A.; Pippel, M.; Young, G. R.; Winkler, S.; Brandl, H.; Henry, I.; Dahl, A.; Powell, S.; Hiller, M.; Myers, E.; Rink, J. C. (2018). “The genome of Schmidtea mediterranea and the evolution of core cellular mechanisms”. Nature. doi:10.1038/nature25473

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An extinct frog that is still living

by Piter Kehoma Boll

Hybrids, as you probably know, are organisms that arise from the mating of two individuals of different species. A mule, for example, is a well known hybrid between a horse and a donkey. Hybrids are usually sterile, although not all of them are, and some of them have a very peculiar way to continue to exist by using a process called hybridogenesis.

Hybrids that rely on hybridogenesis function in the following way: there are two original species, let’s call them A and B. When they copulate with each other, they produce a hybrid offspring, AB, which has half of the genes from one parent and half from the other. In “normal” hybrids, such creatures are completely sterile, unable to produce viable gametes, or can give rise to a new hybrid species by producing mixed gametes. However, in this peculiar kind of hybrids, called kleptons, things work differently.

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Pelophylax kl. hispanicus, the holder of a treasure. Photo by Andreas Thomsen.*

When kleptons are producing gametes, they never recombine the genomes of the two parents, but rather exclude the genome of one of them and produce gametes that contain the genome of the other parent. For example, the hybrid AB produces only A gametes, while the B genome is excluded. This means that if AB mates with a partner of the species A, the offspring will be formed by pure A individuals. If mating with B, the offspring will contain only new AB hybrids.

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The edible frog Pelophylax kl. esculentus is a klepton formed by breeding P. lessonae and P. ridibundus. The klepton only produces gametes of P. ridibundus, eliminating the genome of P. lessonae during meiosis. (Photo by Wikimedia user Darekk2).**

This mode of reproduction is very common in frogs of the genus Pelophylax, as the example seen in the picture above. Another interesting point about kleptons is that they are usually unable to mate with another klepton. They rely one the parent species to reproduce, therefore “parasitizing” them.

A recently published paper on Pelophylax frogs reports a peculiar case in which one of the parent species is extinct. The klepton, known as Pelophylax kl. hispanicus, is the result of P. bergeri crossing with a now extinct species of Pelophylax. The case is that the gametes that P. kl. hispanicus produce are of the extinct species, but they can only fertilize gametes of P. bergeri. In other words, we could say that the extinct species is still alive inside the klepton, relying on P. bergeri to pass to the next generations.

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Pelophylax kl. hispanicus is a klepton that maintains the genome of an extinct species alive. Image extracted from Dubey & Dufresnes (2017).**

The authors suggest that perhaps we could find a way to bring the extinct species back, separated from P. bergeri. Although the result of crossing two P. kl. hispanicus is an sterile offspring, they think that continuous trials may end up revealing an eventual fertile offspring. Is it worth trying? Perhaps. But anyway, this is one more astonishing feature of nature, don’t you agree?

How many more extinct species may be living in a similar way, trapped in a hybrid?

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References:

Wikipedia. Hybridogenesis in water frogs. Available at <https://en.wikipedia.org/wiki/Hybridogenesis_in_water_frogs&gt;. Access on October 12, 2017.

Dubey, S.; Dufresnes, C. (2017) An extinct vertebrate preserved by its living hybridogenetic descendant. Scientific Reports 7: 12768. https://dx.doi.org/10.1038/s41598-017-12942-y

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Friday Fellow: Common Water Bear

by Piter Kehoma Boll

Tiny and tough, our newest Friday Fellow can be found hidden among the moss throughout most of the planet, and perhaps even beyond it, for if there is a species to which space is a piece of cake, that species is the common water bear Milnesium tardigradum.

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A Scaning Electron Microscope (SEM) image of a specimen of the common water bear in its active state. Photo extracted from Schokraie et al. (2012).*

You may have already heard of tardigrades or water bears, tiny chubby animals that are able to withstand the harsher conditions, such as intense desiccation, radiation and even the vacuum of outer space. Most of the data regarding the toughness of these organisms comes from the common water bear, the most widespread species of the phylum Tardigrada.

Measuring up to 0.7 mm in length, the common water bear has eight legs with claws on their end and is considered a predator, feeding on a variety of other small organisms, including algae, rotifers and nematodes. It has a worldwide distribution and is commonly found living on moss, such as the cosmopolitan silvergreen moss already presented here.

As members of the supergroup Ecdysozoa (which also includes arthropods and roundworms), tardigrades undergo ecdysis, also commonly known as molting, a process through which they shed their exoskeleton. In the common water bear, females always lay eggs around the time of molting. Before leaving the old exoskeleton, the females lay the clutch of eggs, which may vary from 1 to 12 eggs, between the old and the new exoskeleton and usually remain inside the old exoskeleton several hours after laying the eggs. When they finally leave, the eggs remain inside the shed skin, which perhaps helps them to be more protected from danger.

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A clutch of seven eggs is left in the empty exoskeleton while the female leaves. Photo by Carolina Biological Supply Company.**

When the habitat of the common water bear gets dry, it enters in a state called cryptobiosis, in which the body shrinks and the metabolism stops. Under this state, known as tun, it can withstand high doses of radiation and both high and zero air pressure, surviving even in the environment of outer space. It is not invincible, however. Radiation in doses above 1000 Gy may not always kill them, but always let them sterile, which is, evolutionary, basically the same thing.

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SEM image of the common water bear in the tun state. Photo extracted from Schokraie et al. (2012).*

Nevertheless, the American cockroach is just an amateur regarding survival when compared to the common water bear.

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References:

Horikawa, D. D.; Sakashita, T.; Katagiri, C.; et al. (2009) Radiation tolerance in the tardigrade Milnesium tardigradumInternatonal Journal of Radiation Biology, 86(12): 843–848. http://dx.doi.org/10.1080/09553000600972956

Schokraie E, Warnken U, Hotz-Wagenblatt A, Grohme MA, Hengherr S, et al. (2012) Comparative proteome analysis of Milnesium tardigradum in early embryonic state versus adults in active and anhydrobiotic state. PLoS ONE 7(9): e45682. https://dx.doi.org/10.1371/journal.pone.0045682

Suzuki, A. C. (2003) Life history of Milnesium tardigradum Doyère (Tardigrada) under a rearing environment. Zoological Science 20(1): 49–57. https://doi.org/10.2108/zsj.20.49

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The bat folk songs: cultural evolution in our winged relatives

by Piter Kehoma Boll

For a long time, culture was considered a human trait, but nowadays we recognize the existence of culture in many other species, such as other primates, whales and some birds too. Now there are some evidences of culture being found in bats too.

A group of researchers from China studied the calls of the Chinese rufous horseshoe bat (Rhinolophus sinicus) across different populations and compared them to genetic and environmental variables to determine whether the differences where linked to genetic differences between the populations or to different environments that would force the bats to change their calls in order to use them more successfully.

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The smile of a cult bat (Rhinolophus sinicus). Photo by Ecohealth Alliance, extracted from Eureka Alert.

The results indicate that none of those two factors were strongly linked to the acoustic differences in the calls. The most likely explanation is that the differences happen due to cultural drift. The bats are teaching a way to speak to their children that is slightly different from what their neighbors speak, even if the neighbors are genetically similar and live in a similar environment.

As an animal’s call is an important variable during mating, this may eventually lead to reproductive isolation even without genetic differences. Culture can also shape evolution!

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Reference:

Xie, L.; Sun, K.; Jiang, T.; Liu, S.; Lu, G.; Jin, L.; Feng, J. (2017) The effects of cultural drift on geographic variation in echolocation calls of the Chinese rufous horseshoe bat (Rhinolophus sinicus)Ethology 123(8): 532-541.

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How do new species form?

by Piter Kehoma Boll

A long, long time ago, I wrote two posts here about the definition of species, explaining briefly the most important horizontal and vertical species concepts. So we all agree that species exists, but how they emerge? How one species become two, or how one species become another?

The phenomenon by which it occurs is called speciation. Well, sort of… It all depends on how you define a species, actually (so be certain to have read the posts I mentioned above).

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Model of a lineage splitting into two lineages that evolve independently and eventually become separated species. Extracted from Hawlitschek et al. (2012)*

Speciation is usually defined as the evolution of reproductive isolation, therefore it deals more with the concept of biological species, but also with the ecological concept and certainly needs some insights on the vertical concepts. If two populations are reproductively isolated, it means that the individuals of one of them are unable or unwilling to breed with those of the other. This usually arrives through genetic and ecological differences that lead to differences in behavior, morphology, physiology. And considering that, we can classify reproductive isolation into two groups: pre-zygotic and post-zygotic isolation.

In pre-zygotic isolation, the two species are reproductively isolated because they do not want or cannot mate and produce an zygote. This may happen simply because of different behaviors in which the two species occupy different places in the environment, mate at different times of the year or even because they are not sexually attracted to each other. There are several experiments using fruitflies that demonstrate how this may evolve pretty fast.

In the late 1980s, William R. Rice and George W. Salt separated individuals of Drosophila melanogaster depending on their preference for dark × light and wet × dry environments, allowing them to mate only with other specimens showing the same preferences. After several generations, the individuals of one group were unable to mate with those of other groups because of their strong habitat preferences, making them unlikely to interact. A similar experiment was performed by Diane Dodd using the species Drosophila pseudoobscura, in which one population was raised with starch as food and other with maltose as food. In this case, after several generations the flies showed a strong preference to mate with individuals of the same group and to reject those of the other group.

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Evolution of reproductive isolation in fruit flies of the species Drosophila pseudoobscura after several generations fed with different sugars.

Such speciation events are called ecological speciation and are also well-documented in the widl, especially regarding fish preferring different habitats, such as shallow × deep water or still × running water. Eventually the individuals will diverge into two groups that are ecologically isolated in the same environment and consequently become reproductively isolated as well.

Post-zygotic isolation is generally a more advanced form of isolation that indicates deep genetic divergences. This is more commonly associated with the notion of biological species and is based on the inability of the individuals of the two species to produce viable offspring. They may mate with each other and even produce a zygote, but this will be unable to developed into an embryo or the offspring will be sterile or otherwise unable to survive enough to breed. A classical example is the mule, the hybrid of a mare and a donkey that is usully sterile.

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A mare, Equus ferus caballus (left), a donkey, Equus africanus asinus (right) and a mule (center). Photos by ‘Little Miss Muffit’ (flickr.com/people/42562654@N00)(mare), Adrian Pingstone (donkey) and Dario Urruty (mule).

In both forms of speciation mentioned above, reproductive isolation usually arises from the accumulation of small differences due to natural selection. This may be enhanced by two phenomena, pleiotropy and genetic hitchhiking.

Pleiotropy is the phenomen by which a single gene have influence over more than one phenotypic trait. For example, a gene that influences the shape of a bird’s bill may also make it change its diet and its song. Several human genetic diseases, such as phenylketonuria (PKU), are examples of pleiotropy.

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The frizzled trait in chickens, which makes the feather curl outward, also leads to delayed sexual maturity and decreased metabolism rate. Photo by flickr user Just chaos.*

Genetic hitchhiking, on the other hand, is the phenomenon by which a gene that is naturally selected carries neighbours genes that are in the same DNA chain with it. In fruitflies, for example, a gene that is linked to courtship behavior may be drawn with the gene linked to a digestive enzyme, so that flies that specialize in one kind of sugar have a different courtship behavior than others specialized in another sugar.

That’s all for now. In a future post, I’ll talk about the geographic and genetic variables in species formation.

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References and further reading:

Bolnick, D. I., Snowberg, L. K., Patenia, C., Stutz, W. E., Ingram, T. & Lau, O. L. 2009. Phenotype-dependent native habitat preference
facilitates divergence between parapatric lake and stream stickleback. Evolution, 63(8): 2004-2016.

Hendry, A. P.2009. Ecological speciation! Or the lack thereof? Canadian Journal of Fisheries and Aquatic Sciences, 66: 1383-1398.

Hoskin, C. J. & Higgie, M. 2010. Speciation via species interactions: the divergence of mating traits within species. Ecology Letters, 13: 409-420.

Maan, M. E., Hofker, K. D., van Alphen, J. J. M. & Seehausen, O. 2006. Sensory drive in cichlid speciation. The American Naturalist, 167(6):
947-954.

Nosil, P. 2008. A century of evolution: Ernst Mayr (1904-2005). Ernst Mayr and the integration of geographic and ecological factors in
speciation. Biological Journal of the Linnean Society, 95: 26-46.

Turelli, M., Barton, N. H. & Coyne, J. A. 2001. Theory and speciation. TRENDS in Ecology and Evolution, 16(7): 330-343.

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