Cospeciation

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Cospeciation and host-parasite associations. From top to bottom:
Cospeciation: host and parasite speciate concurrently
Host switching: speciation as parasite switches hosts and evolves in reproductive isolation
Independent speciation: parasite speciates on same host, reasons unrelated to host
Extinction: parasite goes extinct on host
Missing the boat: host speciates but parasite does not end up reproductively isolated Cospeciation (5 processes) - with key.png
Cospeciation and host-parasite associations. From top to bottom:
Cospeciation: host and parasite speciate concurrently
Host switching: speciation as parasite switches hosts and evolves in reproductive isolation
Independent speciation: parasite speciates on same host, reasons unrelated to host
Extinction: parasite goes extinct on host
Missing the boat: host speciates but parasite does not end up reproductively isolated

Cospeciation is a form of coevolution in which the speciation of one species dictates speciation of another species and is most commonly studied in host-parasite relationships. In the case of a host-parasite relationship, if two hosts of the same species get within close proximity of each other, parasites of the same species from each host are able to move between individuals and mate with the parasites on the other host. [1] However, if a speciation event occurs in the host species, the parasites will no longer be able to "cross over" because the two new host species no longer mate and, if the speciation event is due to a geographic separation, it is very unlikely the two hosts will interact at all with each other. The lack of proximity between the hosts ultimately prevents the populations of parasites from interacting and mating. This can ultimately lead to speciation within the parasite. [2]

Contents

According to Fahrenholz's rule, first proposed by Heinrich Fahrenholz in 1913, when host-parasite cospeciation has occurred, the phylogenies of the host and parasite come to mirror each other. In host-parasite phylogenies, and all species phylogenies for that matter, perfect mirroring is rare. Host-parasite phylogenies can be altered by host switching, extinction, independent speciation, and other ecological events, making cospeciation harder to detect. [3] However, cospeciation is not limited to parasitism, but has been documented in symbiotic relationships like those of gut microbes in primates. [4]

Fahrenholz's rule

In 1913, Heinrich Fahrenholz proposed that the phylogenies of both the host and parasite will eventually become congruent, or mirror each other when cospeciation occurs. [5] More specifically, more closely related parasite species will be found on closely related species of host. Thus, to determine if cospeciation has occurred within a host-parasite relationship, scientists have used comparative analyses on the host and parasite phylogenies.

In 1968, Daniel Janzen proposed an opposing theory to Fahrenholz's rule. Studying cospeciation within plant-insect relationships, he proposed that species have a physiological range of conditions and environments. Over time, conserved traits within a parasitic species allows for survival in a range of conditions or environments. "Ecological fitting", as it is known, means more closely related parasites will share similar traits that pertain to surviving on a particular host. This provides explanation for the congruence of the host-parasite phylogenies. [5] [6]

Parasitic cospeciation

Fahrenholz's rule appears to be observed in the parasitic cospeciation of pocket gophers and chewing lice. [7]

It is seen, too, between Poaceae grasses and Anguininae nematodes, [8] and between some plants and Phyllonorycter leaf-mining moths. [9]

Symbiotic cospeciation

Black smokers provide energy and nutrients to chemoautotrophic bacteria, which in turn have symbiotically cospeciated with deep sea clams. Blacksmoker in Atlantic Ocean.jpg
Black smokers provide energy and nutrients to chemoautotrophic bacteria, which in turn have symbiotically cospeciated with deep sea clams.

Among animals, symbiotic cospeciation is seen between Uroleucon (aphids) and Buchnera (plants in the Orobanchaceae), [10] between deep sea clams and chemoautotrophic bacteria, [11] and between Dendroctonus bark beetles and certain fungi. [12]

Symbiotic cospeciation is found between Crematogaster ants and Macaranga plants, [13] between Ficus fig trees and fig wasps, [14] and between the Poaceae grasses and Epichloe fungi. [15]

False incongruence

The two main hurdles to determining cospeciation using Fahrenholz's rule are instances of false congruence and false incongruence. False congruence occurs when parasite and host phylogenies mirror each other but not due to cospeciation, for instance, if the parasites were to colonize the hosts after the host species had diverged and congruent phylogenies resulted by chance, but this is unlikely. [16] False incongruence, when cospeciation has occurred, but the phylogenies do not mirror each other, is more common, and can be caused by a number of factors; it might also appear to be present if parasites that are present on a host are not detected by the experimenter. [16]

Host switching

Though parasites have been thought to be specialized to a certain host species, it is common for a parasite to colonize a different host that was not previous colonized by the parasite species. If a "host switch" occurs after a cospeciation event, the presence of the parasite on other host species will disrupt any potential congruence in the two phylogenies. Coupled with extinction or independent speciation, phylogenetic comparisons can become complicated and entirely mask the cospeciation event. [17]

Independent speciation

Typically independent speciation does not significantly alter the phylogenetic analysis used to measure cospeciation. However, in combination with extinction, independent speciation can become very problematic when trying to sort out host and parasite phylogenies. Independent speciation occurs when a single population on a single host undergoes speciation resulting in two sister lineages of parasite on a particular host. In other words, the parasite lineage speciates while the host lineage does not. This becomes complicated when the two lineages of parasites then undergo cospeciation with the host. If one of the two parasite lineages goes extinct from the new host lineage, the phylogenies of the host and parasite will begin to break apart. Even though the parasite and host cospeciated together, the phylogenies will not be congruent. [3]

Extinction

After cospeciation, it is possible for a parasite (or symbiont) to become extinct while its host survives. This can happen if, for example, the host species adapts to a new habitat. [3]

"Missing the boat"

Prior to speciation of the hosts, if the distribution of the parasite population among the host population is sporadic, it is possible that when host speciation occurs, it will occur with hosts that do not have the parasite population. This phenomenon is known as "missing the boat". The parasites could potentially cospeciate with their host down the line, however, the parasites could potentially be absent from some host lineages. Like extinction and independent speciation, "missing the boat" alone will probably have a minimal effect on mapping phylogenies, however, in conjunction with independent speciation, parasite and host phylogenies can begin to break apart. [3]

Related Research Articles

<span class="mw-page-title-main">Parasitism</span> Relationship between species where one organism lives on or in another organism, causing it harm

Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson characterised parasites as "predators that eat prey in units of less than one". Parasites include single-celled protozoans such as the agents of malaria, sleeping sickness, and amoebic dysentery; animals such as hookworms, lice, mosquitoes, and vampire bats; fungi such as honey fungus and the agents of ringworm; and plants such as mistletoe, dodder, and the broomrapes.

<span class="mw-page-title-main">Fig wasp</span> Group of mostly pollinating insects whose larvae live in figs

Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside figs. Some are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, while the pollinators are in the family Agaonidae. While pollinating fig wasps are gall-makers, the remaining types either make their own galls or usurp the galls of other fig wasps; reports of their being parasitoids are considered dubious.

<span class="mw-page-title-main">Louse</span> Order of insects

Louse is the common name for any member of the clade Phthiraptera, which contains nearly 5,000 species of wingless parasitic insects. Phthiraptera has variously been recognized as an order, infraorder, or a parvorder, as a result of developments in phylogenetic research.

<span class="mw-page-title-main">Coevolution</span> Two or more species influencing each others evolution

In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well as gene-culture coevolution.

Coextinction and cothreatened refer to the phenomenon of the loss or decline of a host species resulting in the loss or endangerment of another species that depends on it, potentially leading to cascading effects across trophic levels. The term was originated by the authors Stork and Lyal (1993) and was originally used to explain the extinction of parasitic insects following the loss of their specific hosts. The term is now used to describe the loss of any interacting species, including competition with their counterpart, and specialist herbivores with their food source. Coextinction is especially common when a keystone species goes extinct.

The Prodoxidae are a family of moths, generally small in size and nondescript in appearance. They include species of moderate pest status, such as the currant shoot borer, and others of considerable ecological and evolutionary interest, such as various species of "yucca moths".

<span class="mw-page-title-main">Roderic D. M. Page</span> New Zealand-born evolutionary biologist

Roderic Dugald Morton Page is a New Zealand-born evolutionary biologist at the University of Glasgow, Scotland, and the author of several books. As of 2015 he is professor at the University of Glasgow and was editor of the journal Systematic Biology until the end of 2007. His main interests are in phylogenetics, evolutionary biology and bioinformatics.

The Red Queen's hypothesis is a hypothesis in evolutionary biology proposed in 1973, that species must constantly adapt, evolve, and proliferate in order to survive while pitted against ever-evolving opposing species. The hypothesis was intended to explain the constant (age-independent) extinction probability as observed in the paleontological record caused by co-evolution between competing species; however, it has also been suggested that the Red Queen hypothesis explains the advantage of sexual reproduction at the level of individuals, and the positive correlation between speciation and extinction rates in most higher taxa.

Avian malaria is a parasitic disease of birds, caused by parasite species belonging to the genera Plasmodium and Hemoproteus. The disease is transmitted by a dipteran vector including mosquitoes in the case of Plasmodium parasites and biting midges for Hemoproteus. The range of symptoms and effects of the parasite on its bird hosts is very wide, from asymptomatic cases to drastic population declines due to the disease, as is the case of the Hawaiian honeycreepers. The diversity of parasites is large, as it is estimated that there are approximately as many parasites as there are species of hosts. As research on human malaria parasites became difficult, Dr. Ross studied avian malaria parasites. Co-speciation and host switching events have contributed to the broad range of hosts that these parasites can infect, causing avian malaria to be a widespread global disease, found everywhere except Antarctica.

<span class="mw-page-title-main">Entorrhizomycetes</span> Class of fungi

Entorrhizomycetes is the sole class in the phylum Entorrhizomycota, within the Fungi subkingdom Dikarya along with Basidiomycota and Ascomycota. It contains three genera and is a small group of teliosporic root parasites that form galls on plants in the Juncaceae (rush) and Cyperaceae (sedge) families. Prior to 2015 this phylum was placed under the subdivision Ustilaginomycotina. A 2015 study did a "comprehensive five-gene analyses" of Entorrhiza and concluded that the former class Entorrhizomycetes is possibly either a close sister group to the rest of Dikarya or Basidiomycota.

Host–parasite coevolution is a special case of coevolution, where a host and a parasite continually adapt to each other. This can create an evolutionary arms race between them. A more benign possibility is of an evolutionary trade-off between transmission and virulence in the parasite, as if it kills its host too quickly, the parasite will not be able to reproduce either. Another theory, the Red Queen hypothesis, proposes that since both host and parasite have to keep on evolving to keep up with each other, and since sexual reproduction continually creates new combinations of genes, parasitism favours sexual reproduction in the host.

<span class="mw-page-title-main">Ecological fitting</span> Biological process

Ecological fitting is "the process whereby organisms colonize and persist in novel environments, use novel resources or form novel associations with other species as a result of the suites of traits that they carry at the time they encounter the novel condition". It can be understood as a situation in which a species' interactions with its biotic and abiotic environment seem to indicate a history of coevolution, when in actuality the relevant traits evolved in response to a different set of biotic and abiotic conditions.

Incomplete lineage sorting, also termed hemiplasy, deep coalescence, retention of ancestral polymorphism, or trans-species polymorphism, describes a phenomenon in population genetics when ancestral gene copies fail to coalesce into a common ancestral copy until deeper than previous speciation events. It is caused by lineage sorting of genetic polymorphisms that were retained across successive nodes in the species tree. In other words, the tree produced by a single gene differs from the population or species level tree, producing a discordant tree. Whatever the mechanism, the result is that a generated species level tree may differ depending on the selected genes used for assessment. This is in contrast to complete lineage sorting, where the tree produced by the gene is the same as the population or species level tree. Both are common results in phylogenetic analysis, although it depends on the gene, organism, and sampling technique.

<span class="mw-page-title-main">Eichler's rule</span> Coevolutionary principle regerding parasites and hosts

Eichler's rule is one of several coevolutionary rules which states that parasites tend to be highly specific to their hosts, and thus it seems reasonable to expect a positive co-variation between the taxonomic richness of hosts and that of their parasites.

<span class="mw-page-title-main">Escape and radiate coevolution</span>

Escape and radiate coevolution is a hypothesis proposing that a coevolutionary 'arms-race' between primary producers and their consumers contributes to the diversification of species by accelerating speciation rates. The hypothesized process involves the evolution of novel defenses in the host, allowing it to "escape" and then "radiate" into differing species.

<span class="mw-page-title-main">Ecological speciation</span>

Ecological speciation is a form of speciation arising from reproductive isolation that occurs due to an ecological factor that reduces or eliminates gene flow between two populations of a species. Ecological factors can include changes in the environmental conditions in which a species experiences, such as behavioral changes involving predation, predator avoidance, pollinator attraction, and foraging; as well as changes in mate choice due to sexual selection or communication systems. Ecologically-driven reproductive isolation under divergent natural selection leads to the formation of new species. This has been documented in many cases in nature and has been a major focus of research on speciation for the past few decades.

<span class="mw-page-title-main">Biological rules</span> Generalized principle to describe patterns observed in living organisms

A biological rule or biological law is a generalized law, principle, or rule of thumb formulated to describe patterns observed in living organisms. Biological rules and laws are often developed as succinct, broadly applicable ways to explain complex phenomena or salient observations about the ecology and biogeographical distributions of plant and animal species around the world, though they have been proposed for or extended to all types of organisms. Many of these regularities of ecology and biogeography are named after the biologists who first described them.

Reproductive coevolution in <i>Ficus</i>

The genus Ficus is composed of 800 species of vines, shrubs, and trees, defined by their syconiums, the fruit-like vessels that either hold female flowers or pollen on the inside. In addition to being cultivated by humans for thousands of years, Ficus is also known for their reproductive mutualism with the fig wasp.

<span class="mw-page-title-main">Harrison's rule</span>

Harrison's rule is an observation in evolutionary biology by Launcelot Harrison which states that in comparisons across closely related species, host and parasite body sizes tend to covary positively.

<span class="mw-page-title-main">Phylogenetic reconciliation</span> Technique in evolutionary study

In phylogenetics, reconciliation is an approach to connect the history of two or more coevolving biological entities. The general idea of reconciliation is that a phylogenetic tree representing the evolution of an entity can be drawn within another phylogenetic tree representing an encompassing entity to reveal their interdependence and the evolutionary events that have marked their shared history. The development of reconciliation approaches started in the 1980s, mainly to depict the coevolution of a gene and a genome, and of a host and a symbiont, which can be mutualist, commensalist or parasitic. It has also been used for example to detect horizontal gene transfer, or understand the dynamics of genome evolution.

References

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