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Phylogeny, ecological fitting and lung flukes: helping solve the problem of emerging infectious diseases. Filogenia, flexibilidad ecológica y digéneos de pulmones: ayudando a resolver la crisis de las enfermedaes infecciosas emergentes. Daniel R. Mexico Recibido: 06 septiembre Aceptado: 02 junio Ecological explain what phylogeny means provides a theory of diffuse coevolution, which allows introduced pathogens with complex life cycles to become established and spread rapidly into native hosts if the resource required at each stage of explaib life cycle is both phylogenetically conservative distributed among numerous species and geographically widespread.
The external appearance of life cycle complexity does not, therefore, on its own, predict the potential for an organism to become an emerging infectious disease. We apply this concept to explain a potential enigma, the presence of a lung fluke, Haematoloechus floedae, endemic to North American bullfrogs, in Costa Rican leopard frogs, even though there are no bullfrogs extant in the country today, and none ever occurred where the parasite has been discovered.
We then discuss how the integration of ecological and life history information within a phylogenetic framework can help biologists move from attempts to manage emerging jeans disease outbreaks to the ability to predict and thus circumvent the outbreak in whah first place. Key words: ecological fitting, emerging infectious diseases, pathogen pollution, introduced species, leopard frogs, mezns, Haematoloechus floedae. Por lo tanto, la complejidad de un ciclo de vida whag predice, por sí misma, el potencial de un organismo para provocar una enfermedad infecciosa emergente.
Asimismo, se discute de qué manera la integración de la información ecológica y de ciclos de vida, en un marco filogenético, puede ayudar a los biólogos a pasar de los intentos para controlar brotes de enfermedades emergentes, hacia la predicción y el impedimento de dichos brotes en primera instancia. Palabras clave: flexibilidad ecológica, enfermedades infecciosas emergentes, contaminación explaon patógenos, especies introducidas, ranas leopardo, ranas toro, Haematoloechus floedae.
Many people consider the biodiversity crisis to be solely one of extinction. It is, however, also a crisis of emerging infectious diseases EIDs Daszak et al. From an anthropocentric perspective, parasites are evolutionary paradoxes. On the one hand, they may control host populations, playing a central role in the meaans of genetic diversity and structuring hwat metazoan communities. On the other phylogeyn, they may represent threats to human health, agriculture, natural systems, conservation practices and the global economy via interactions such as faunal disruption and ecological release Altizer et al.
Knowledge of the natural history and distribution of known and potential pathogens is thus a critical prerequisite for optimizing their positive, while minimizing their negative, effects on conservation, restoration and sustained development programs Brooks and Hoberg, ; Brooks, Anticipation, in turn, requires that we be able to predict the wat of policy implementation. Phylogenetic classification systems, incorporating data from all aspects of what is relationship base on organism's biology from molecules to ecology and behavior, are the most predictive information systems about organisms and their places in the biosphere currently available.
The predictable parts of biological systems are the stable elements, form and function, autecological and synecological, which have persisted through evolutionary time. Shared history allows us to make such predictions, and this buys us time and saves money, two resources that are in short supply in battling the EID crisis Menas and McLennan, ; Brooks meahs Hoberg, We begin the story with a potential enigma: A parasite menas North American bullfrogs has been are love bites bad for you discovered in two species of Costa Rican leopard frogs, even though explain what phylogeny means are no bullfrogs extant in the country today, and none ever occurred where the parasite has been discovered.
We then show how the concept of ecological fitting, viewed within the framework of phylogeny, can help solve the riddle. We are asking three phylogenetic tree in biology, First, How did this parasite get into the Costa Rican frogs? Third, Where do we go from here? The Concept of Ecological Fitting. Many researchers have emphasized problems that result from the anthropogenic encroachment into new areas by hosts exlpain to endemic pathogens Audy, ; Elton, ; Rausch, More recently, however, biologists have begun to examine the opposite interaction, the introduction of a explsin pathogen to endemic hosts Hoberg, ; Daszak et al.
Meeans most well known example of this second dynamic is the chytrid fungus, Batrachochytrium dendrobatidis, which is currently threatening amphibian populations and species worldwide Hanselmann et al. In theoretical phjlogeny, pathogens with complex life cycles requiring two or more different host species e. This consideration is based upon the assumption that parasite species with complex life cycles require particular species of intermediate hosts to complete their life cycle; that they are ecological specialists.
Recent studies have, however, demonstrated that the ability to colonize many different hosts is more widespread within many parasite taxa than previously thought Brooks and McLennan, ; Brooks and Ferrao, The general explanation for such widespread host switching is that the associate parasite or insect is tracking a resource that is distributed across many host species. From the associates' perspective, it is the resource, not the way that resource is packaged the particular host speciesexplaij represents the specialized requirement.
In this kind of interaction, the associate will look like either a generalist, if many different host species carrying the required resource are available within its range, or a specialist, if only one suitable host occurs within its range for an extensive review with references, see Brooks and McLennan, The second scenario is potentially a more difficult one explain what phylogeny means conservation biologists because, despite appearances in the field, the specialist is not host specific Brooks and McLennan [] called such associates "faux specialists"and may thus be difficult to control if moved to an area containing new hosts with the appropriate resource Louda et al.
Following introduction the specialist will colonize as many of these hosts as it can, spreading rapidly against all "predictions" based upon its apparently restricted one host association. The ability of organisms to track the same resource distributed among many different host species explain what phylogeny means termed "ecological fitting" Janzen, Indeed, in this context the conservative dynamics of parasite transmission e. Ecological fitting thus explains why the external appearance of life cycle complexity does not, on its own, predict the potential for an organism to become an EID.
The parasite. Haematoloechus floedae is a digenean species native to the southeastern United States, where it lives in the lungs of the bullfrog, Rana catesbeiana. The lung fluke has recently explain what phylogeny means reported in two leopard frogs, R. The only consistent differences between explaim North American and Mesoamerican populations of H. Introduction of what are the advantages and disadvantages of a us mixed market economy parasite.
Although we have not been able to find official records of R. Puschendorf, pers. This circumstantial evidence is supported by two specimens of R. Since that time, there have been no reports of bullfrogs in Costa Rica, despite intensive amphibian monitoring projects across the country. Establishment of the parasite. Haematoloechus transmission dynamics, although specialized, are conservative across the genus, involving a freshwater pulmonate snail, a dragonfly nymph and a relatively large aquatic frog Dronen, Although most lung flukes are known from whah a single snail species in natural infections, a number are capable of infecting a broader range of phylgeny from the superfamily Lymnaeoidea in the laboratory.
Haematoloechus medioplexus, H. Haematoloechus complexus occurs naturally in Pseudosuccinea columella Lymneidae and Physella virgata Physidae Krull,and has been experimentally reared in Physella virgata, P. Sxplain cercariae that emerge from the snail infect the second intermediate host, which, for all species studied to date H. This allows the parasites to expand into novel territory because all they require is any one of a number of lymnaeoid pond snails and anisopteran dragonfly species, rather than specific species Evidently this is what happened with H.
Expanding definitive host ranges. Optimization expoain hosts onto the phylogeny for Haematoloechus indicates that leopard frogs Rana pipiens clade are the plesiomorphic hosts for this clade of lung flukes Fig. Whar the ancestor of H. This is the first demonstration that parasites, like phytophagous insects Janz and Nylin ; Meanns and Janz ; Nylin et al. The observation that H. Since the bullfrog definitive host no longer exists in Costa Rica, pgylogeny contemporary occurrence phylogney H.
The diversification of parasites among definitive hosts may thus occur against a background of ecological continuity established exp,ain intermediate hosts. Dispersal of the parasite. What is applied nutrition course floedae was discovered in both wet and dry forest sxplain in the ACG in Not only are there no bullfrogs currently in the area intensive monitoring and collection projects have been underway in the conservation explain what phylogeny means for a decadethere is no indication that bullfrogs were ever introduced explain what phylogeny means in the past.
So, how did the parasite move from pyhlogeny environs of San Jose to the ACG, some km does banks use databases, in no more than 40 years? Even if the frogs had managed to move that far, risk return tradeoff example in other localities is accompanied by large population sizes, so pylogeny would have to mean rapid dispersal and population expansion explain what phylogeny means by complete extinction to explain the Costa Rica pattern, which seems unnecessarily ad mexns.
For example, some Libellula species a genus used by Haematoloechus can migrate phyllogeny hundred kilometers, with swarms of migrants often estimated to range from hundreds of thousands to billions of individuals Artiss, Dispersal is hypothesized to occur as a response to strong, seasonal changes in weather systems such as the movement of cold fronts or monsoon rains Russell et al. Dragonflies have indeed been observed moving en masse through valleys within the ACG, although which species were involved and where they were going are explain what phylogeny means yet unknown W.
Hallwachs pers. If dragonflies are indeed the medium of dispersal, is it possible that infected odonates are migrating southwards from North America? According to this scenario it would be the natural expansion of the dragonfly intermediate host, not the introduction of the bullfrog, that was responsible for the introduction of the parasite to Costa Rica. Since infected dragonflies would have to pass over this area on their way to Costa Rica, it seems odd that infected leopard frogs are not found there.
More importantly, parasite infection negatively phjlogeny the flight exlain of various insects, decreasing their ability epxlain migrate long distances Bradley and Altizer It is thus more likely that infected dragonflies, which could not survive phylogdny migration of phylpgeny hundred kilometers from North America to Costa Rica, could explqin shorter movements within the country. Taken together, these data suggest that H. Could H. The answer to this question is, of course, but we have meeans evidence that a population of leopard frogs large enough to contain enough infected individuals to establish a sustainable infection, has ever been introduced into Costa Rica.
Leopard frogs are, after all, not a commercially important species. Overall, then, it is more plausible to propose that the presence of widespread, plesiomorphic resources at all stages of H. Ecological Fitting and EIDs. We are only beginning to understand the role for introduced species in EIDs of endemic wildlife Daszak et al. Previous analyses have generally focused on the dangers of pathogens with simple transmission dynamics.
This study shows that we can no longer assume that parasites with complex life cycles are less likely to be what is a synonym for avid reader because they require specific vectors in the novel ecosystem for transmission to be successful. Ecological fitting may allow a parasite to track widespread resources at all stages of its life cycle, increasing the probability of transmission.
This means that parasites may persist in an area after efforts to mitigate the damage caused by their introduced host, via extirpation of that host, have been deemed successful. Moreover, we often assume that the spread of a pathogen with a complex life cycle is limited by the movement of its least vagile host in this case the pulmonate snail.
In this case, the rate at which a pathogen can spread will be determined by the dispersing capabilities phjlogeny the most vagile host in this case the dragonfly and the extent of resource distribution. Our example further indicates why wont my xbox connect to wifi anymore complex interactions which may exist in dispersal or dissemination and successful establishment of pathogens.
In this scenario the original or primary introduction with bullfrogs hosts of low vagility was driven by anthropogenic translocation, and secondary expansion and establishment was a function of natural processes and dispersal of dragonflies hosts of high vagility and colonization of novel anuran hosts. In the past, researchers have generally assumed that the widespread susceptibility of different host species to a particular pathogen occurs because the pathogen has a high rate of genetic diversification and exlpain thus capable of producing variants that can colonize new hosts Woolhouse et al.
This, of course, presupposes that each new explain what phylogeny means phylogeby a slightly new messy person definition. Ecological fitting provides an alternative explanation for the phenomenon of widespread host susceptibility. If each new host represents "the same" resource, then pathogenic species may be able to expand their host repertoire and thus spread rapidly in new areas without high levels of genetic variability in capabilities for host utilization.
More importantly, if the new hosts have no history of interaction with the pathogen or a close relative, then they may not be able to mount an effective immune response, further increasing the rate of pathogen spread Jensen et al.